Trends in Diporeia populations across the Laurentian Great Lakes, 1997-2009

Benthic communities in the Laurentian Great Lakes have been in a state of flux since the arrival of dreissenid mussels, with the most dramatic changes occurring in population densities of the amphipod Diporeia. In response, the US EPA initiated an annual benthic macroinvertebrate monitoring program on all five Great Lakes in 1997. Although historically the dominant benthic invertebrate in all the lakes, no Diporeia have been found in Lake Erie during the first 13 years of our study, confirming that Diporeia is now effectively absent from that lake. Populations have almost entirely disappeared from our shallow (< 90 m) sites in lakes Ontario, Huron, and Michigan. In Lake Ontario, three of our four deep (> 90 m) sites still supported Diporeia populations in 2009, with densities at those sites ranging between 96 and 198/m². In Lake Michigan, populations were still found at six of our seven deep sites in 2009, with densities ranging from 57 to 1409/m². Densities of Diporeia in 2009 at the four deep sites in Lake Huron were somewhat lower than those in Lake Michigan, ranging from 191 to 720/m². Interannual changes in population size in Lake Huron and Lake Michigan have shown a degree of synchrony across most sites, with periods of rapid decline (1997-2000, 2003-2004) alternating with periods of little change or even increase (2001-2002, 2005-2009). There has been no evidence of directional trends at any sites in Lake Superior, although substantial interannual variability was seen.     

Evidence for Remote Effects of Dreissenid Mussels on the Amphipod Diporeia: Analysis of Lake Ontario Benthic Surveys, 1972–2003

The status of invasive dreissenid mussels (Dreissena polymorpha and D. bugensis) and native amphipods (Diporeia spp.) in Lake Ontario was assessed in 2003 and compared with historical data. D. polymorpha (zebra mussels) were rarely observed in 2003, having been displaced by D. bugensis (quagga mussels). D. bugensis expanded its depth range from 38 m depth in 1995 to 174 m in 2003 and this dreissenid reached densities averaging 8,000/m² at all sites < 90 m. During the same time period, Diporeia populations almost completely disappeared from 0–90 m depth, continuing a declining trend from 1994–1997 reported in previous studies. The average density of Diporeia in the 30–90 m depth interval decreased from 1,380/m² to 63/m² between 1997 and 2003. Prior to 2003, areas deeper than 90 m represented a refuge for Diporeia, but even these deep populations decreased, with densities declining from 2,181/m2 in 1999 to 545/m2 in 2003. Two common hypotheses for the decline of Diporeia in the Great Lakes are food limitation and a toxin/pathogen associated with dreissenid pseudofeces. The Diporeia decline in deep waters preceded the expansion of D. bugensis to these depths, and suggests that shallow dreissenid populations remotely influence profundal habitats. This pattern of decline is consistent with mechanisms that act from some distance including nearshore dreissenid grazing and downslope transport of pseudofeces.     

Investigation of viruses in Diporeia spp. from the Laurentian Great Lakes and Owasco Lake as potential stressors of declining populations

The benthic amphipod Diporeia is an ecologically and biogeochemically important constituent of deep freshwater lakes in North America. The proliferation of dreissenid mussels in the mid-1990s coincided with a sharp decrease in Diporeia populations in several Laurentian Great Lakes; however the ultimate cause and mechanisms of their decline are still unknown. Here we examined the composition of DNA viruses associated with Diporeia collected from populations of Lake Michigan that had declined and stable populations in Lake Superior and Owasco Lake (Finger Lake in central New York State). Viral metagenomic libraries from Owasco Lake and Lake Superior were comprised primarily of bacteriophages, which may infect bacteria within the amphipod microbiome. In contrast, the metagenomic library from Lake Michigan contained well-represented ssDNA circular viral genomes. The prevalence and viral load of one putative Type V ssDNA circular virus (LM29173) that recruited almost 30% of total viral sequence reads in the Lake Michigan library was analyzed by quantitative PCR. The prevalence of LM29173 was over two orders of magnitude greater in Lake Michigan compared to the other two lakes. Although further research is necessary to establish the pathology and epidemiological extent of viral-Diporeia interactions, our data suggest that viruses may be numerically significant constituents of the Diporeia microbiome, and if pathogenic some of these viruses may be a stressor of Great Lakes Diporeia populations. Our data further indicate that special attention should be given to the circovirus that was prevalent in the declining Michigan population but uncommon in the other two lakes.     

Fall diet and bathymetric distribution of deepwater sculpin (Myoxocephalus thompsonii) in Lake Huron

Deepwater sculpin Myoxocephalus thompsonii are an important component of Great Lake's offshore benthic food webs. Recent declines in deepwater sculpin abundance and changes in bathymetric distribution may be associated with changes in the deepwater food web of Lake Huron, particularly, decreased abundance of benthic invertebrates such as Diporeia. To assess how deepwater sculpins have responded to recent changes, we examined a fifteen-year time series of spatial and temporal patterns in abundance as well as the diets of fish collected in bottom trawls during fall of 2003, 2004, and 2005. During 1992–2007, deepwater sculpin abundance declined on a lake-wide scale but the decline in abundance at shallower depths and in the southern portion of Lake Huron was more pronounced. Of the 534 fish examined for diet analysis, 97% had food in the stomach. Mysis, Diporeia, and Chironomidae were consumed frequently, while sphaerid clams, ostracods, fish eggs, and small fish were found in only low numbers. We found an inverse relationship between prevalence of Mysis and Diporeia in diets that reflected geographic and temporal trends in abundance of these invertebrates in Lake Huron. Because deepwater sculpins are an important trophic link in offshore benthic food webs, declines in population abundance and changes in distribution may cascade throughout the food web and impede fish community restoration goals.     

Trophic connections in Lake Superior Part II: The nearshore fish community

We use detailed diet analyses of the predominant planktivorous, benthivorous and piscivorous fish species from Lake Superior to create a nearshore (bathymetric depths < 80 m) fish community food web. The food web was based on analysis of 5125 fish stomachs collected seasonally (spring, summer, fall) from 9 nearshore sites in 2005. Based on mass of prey items, nearshore diets across all sites and seasons were similarly structured with a dominance of macroinvertebrates (Mysis diluviana and Diporeia spp). Although the piscivorous fishes like lean lake trout (Salvelinus namaycush) fed to a lesser extent on Diporeia and Mysis, they were still strongly connected to these macroinvertebrates, which were consumed by their primary prey species (sculpin spp., rainbow smelt Osmerus mordax, and coregonines). The addition of Bythotrephes to summer/fall cisco and lake whitefish diets, and the decrease in rainbow smelt in lean lake trout diets (replaced by coregonines) were the largest observed differences relative to historic Lake Superior diet studies. Although the offshore food web of Lake Superior was simpler than nearshore in terms of number of fish species present, the two areas had remarkably similar food web structures, and both fish communities were primarily supported by Mysis and Diporeia. We conclude that declines in Mysis or Diporeia populations would have a significant impact on energy flow in Lake Superior. The food web information we generated can be used to better identify management strategies for Lake Superior.     

Trophic connections in Lake Superior Part I: The offshore fish community

Detailed diet linkages within the offshore (> 80 m bathymetric depth) food web of Lake Superior are currently not well identified. We used analyses of fish stomach contents to create an empirically based food web model of the Lake Superior offshore fish community. Stomachs were collected seasonally (spring, summer, and fall) from nine offshore locations in 2005, using bottom and midwater trawls. In total, 2643 stomachs representing 12 fish species were examined. The predominant fish species collected were deepwater sculpin (Myoxocephalus thompsonii), siscowet (Salvelinus namaycush siscowet), kiyi (Coregonus kiyi), and cisco (Coregonus artedi). Mysis diluviana was the most common prey item, indicating that changes in Mysis abundance could have a profound impact on the entire offshore food web. Mysis was the primary diet item of deepwater sculpin (≥ 53% by mass) and kiyi (≥ 96% by mass) regardless of depth or season. The invasive Bythotrephes was an important diet component of the pelagic cisco in summer and fall. Deepwater sculpin were the primary diet item of siscowet (≥ 52% by mass), with coregonines appearing in the diet of larger (> 400 mm) siscowet. Non-metric multidimensional scaling analysis indicated that there were no statistically significant seasonal or site-specific differences in diets of deepwater sculpin, cisco, or kiyi. Site was the primary structuring factor in siscowet diets. Generally, in Lake Superior, the diet items of the dominant offshore species did not appear to be in danger from those types of major ecological shifts occurring in the lower Laurentian Great Lakes.     

Prey selection by the Lake Superior fish community

Mysis diluviana is an important prey item to the Lake Superior fish community as found through a recent diet study. We further evaluated this by relating the quantity of prey found in fish diets to the quantity of prey available to fish, providing insight into feeding behavior and prey preferences. We describe the seasonal prey selection of major fish species collected across 18 stations in Lake Superior in spring, summer, and fall of 2005. Of the major nearshore fish species, bloater (Coregonus hoyi), rainbow smelt (Osmerus mordax), and lake whitefish (Coregonus clupeaformis) consumed Mysis, and strongly selected Mysis over other prey items each season. However, lake whitefish also selected Bythotrephes in the fall when Bythotrephes were numerous. Cisco (Coregonus artedi), a major nearshore and offshore species, fed largely on calanoid copepods, and selected calanoid copepods (spring) and Bythotrephes (summer and fall). Cisco also targeted prey similarly across bathymetric depths. Other major offshore fish species such as kiyi (Coregonus kiyi) and deepwater sculpin (Myoxocephalus thompsoni) fed largely on Mysis, with kiyi targeting Mysis exclusively while deepwater sculpin did not prefer any single prey organism. The major offshore predator siscowet lake trout (Salvelinus namaycush siscowet) consumed deepwater sculpin and coregonines, but selected deepwater sculpin and Mysis each season, with juveniles having a higher selection for Mysis than adults. Our results suggest that Mysis is not only a commonly consumed prey item, but a highly preferred prey item for pelagic, benthic, and piscivorous fishes in nearshore and offshore waters of Lake Superior.     

Spatial variation in RNA:DNA ratios of Diporeia spp. in the Great Lakes region

Over the past two decades, Diporeia in all of the Laurentian Great Lakes, except Superior, have declined dramatically. These declines have seemingly coincided with expansion of invasive Dreissena polymorpha and D. bugensis, however the exact mechanisms underlying decreasing Diporeia densities are obscure. We explored the use of RNA:DNA (R/D) ratios as a conditional index for Diporeia by experimentally demonstrating that Diporeia R/D responds to periods of starvation. Moreover, during 2008–2009 we collected Diporeia from throughout the Great Lakes and Cayuga Lake (New York, USA), and used R/D ratios to index condition of these in situ collected animals. We evaluated spatial and temporal variation of nucleic acid indices using classification and regression tree (CART) analysis with a suite of environmental variables included as potential predictors. Diporeia R/D of in situ collected specimens exhibited pronounced spatial and temporal variation, but multiple CART models described only a small amount of this variation. While we observed some variation in Diporeia R/D among lakes, nucleic acid ratios appeared to respond weakly to Diporeia population characteristics and local environmental conditions. Specifically, CART analyses revealed that Diporeia R/D was particularly low at extreme depths, and interestingly, Diporeia nucleic acids were not strongly associated with the presence of dreissenids. In summary, while a limited amount of variation in Diporeia R/D was attributable to environmental conditions, the majority of Diporeia R/D variation was unaccounted for. Hence, the causative factors underlying spatio-temporal variation of Diporeia R/D and the mechanistic reasons for Diporeia declines in the Great Lakes remain largely unknown.     

Coexistence of the native benthic amphipod Diporeia spp. and exotic dreissenid mussels in the New York Finger Lakes

Populations of the benthic amphipod Diporeia spp. have sharply declined since the early 1990s in all North America's Great Lakes except Lake Superior. The onset and continued decline coincides with the invasion of these lakes by zebra (Dreissena polymorpha) and quagga (Dreissena rostriformis bugensis) mussels and the spread of quagga mussels to deep habitats. The six deepest Finger Lakes of central New York (Seneca, Cayuga, Skaneateles, Canandaigua, Keuka, and Owasco) have historically been Diporeia habitat and have had dreissenids for more than a decade. These lakes represent a wide range of trophic state, maximum depth, and dreissenid invasion history. We hypothesized that Diporeia abundance would be negatively impacted by dreissenid mussel expansion in the Finger Lakes. During 2006–2010, we sampled Diporeia and mussel populations in these six lakes. Diporeia was present in all six lakes, and was abundant (2000/m²) in Owasco Lake that has only zebra mussels and in Cayuga and Seneca Lakes that have had zebra and quagga mussels since 1994. Diporeia abundance was lowest (1000/m²) in Skaneateles, Canandaigua, and Keuka Lakes where quagga mussels have recently expanded. Productivity indicators explained much of the variability of Diporeia abundance. The persistence of Diporeia with quagga mussels in these lakes may be because of available alternative food resources. Fatty acid tracers indicate that Diporeia from Owasco Lake, the lake without quagga mussels, utilize diatoms, but Diporeia from Cayuga Lake that coexist with abundant quagga mussels also use food resources associated with terrestrial detritus that cannot be intercepted by dreissenids.     

The Quagga Mussel Invades the Lake Superior Basin

Prior studies recognized the presence of a single dreissenid species in Lake Superior—the zebra mussel Dreissena polymorpha. However, taxonomic keys based on traditional shell morphology are not always able to differentiate dreissenid species with confidence. We thus employed genetic and morphological analyses to identify dreissenids in a major river-embayment of Lake Superior—the lower St. Louis River/Duluth-Superior Harbor—during 2005–2006. Our results revealed the presence of a second dreissenid species—the quagga mussel D. bugensis (alternatively known as D. rostriformis bugensis). Both species occurred in mixed clusters, in which zebra mussels outnumbered quagga mussels (20–160:1). The largest quagga mussel collected in 2005 was 26.5 mm long and estimated to be two years old, suggesting that the initial introduction occurred no later than 2003. Further monitoring is necessary to determine whether the quagga mussel will colonize Lake Superior. Our results indicate that the coupling of conventional morphological and molecular approaches is essential for monitoring dreissenid species.     

Movement and growth indicators in resident and adfluvial coaster brook trout (Salvelinus fontinalis) in the Hurricane River,Lake Superior,Michigan, USA

Brook trout (Salvelinus fontinalis) are found throughout Lake Superior, Lake Nipigon, and their tributaries. Lacustrine and adfluvial life history variants were historically popular with anglers and were called coasters; coaster brook trout populations are now severely reduced and are of conservation concern. Coasters were known to grow larger and mature later than their stream resident counterparts. This study compared movement patterns, age, size, condition, and relative weight of wild coaster and resident brook trout from the Hurricane River, Pictured Rocks National Lakeshore, Michigan. Wild brook trout ≥ 100 mm from the Hurricane River downstream from Hurricane Falls were tagged with passive integrated transponder tags and monitored for stream-lake movement behavior from May 2003 to November 2007. During 2006 and 2007, brook trout were scale sampled and aged to construct a regression that was then used to calculate the age of all brook trout tagged from 2003 to 2007. Most brook trout movement took place in the fall with October the peak month of emigration with a secondary peak in late spring/early summer and some activity nearly year round. There were no differences found in age structure, size or condition between coasters and residents while in the stream. Our data suggest that a priori growth differences are not determining the expression of coaster outmigration and that stream-lake movements made by coasters, likely driven by habitat requirements, may be highly flexible and facultative.     

Diet effects on lipid composition,somatic growth potential,and survival of the benthic amphipod Diporeia spp.

The benthic amphipod Diporeia represents a crucial trophic link that conveys vital nutrients and energy to predators at higher trophic levels. The current decline of Diporeia populations, mostly in the North American Great Lakes, may, in part, be related to concurrent declines in food quantity and/or quality. We hypothesized that somatic growth and survival of Diporeia would be positively related to dietary supply and subsequent retention of polyunsaturated fatty acids (PUFA); a class of chemicals known to affect diet quality. We examined how different algal PUFA concentrations in; a) Ankistrodesmus falcatus (Chlorophyta), b) a naturally occurring diatom assemblage from Lake Ontario, c) a non-toxic strain of Microcystis aeruginosa (Cyanophyta), and, d) fasting for 30 d, affected PUFA concentrations, somatic growth, and survival of Diporeia. Total PUFA concentrations were significantly higher in A. falcatus than in diatoms and Microcystis, but only diatoms contained considerable amounts of eicosapentaenoic (EPA) and docosahexaenoic acid (DHA). EPA, DHA, and arachidonic acid (ARA) were highly retained in Diporeia even in the absence of dietary supply with ARA being the most efficiently bioaccumulated PUFA. Survivorship of Diporeia ranged from 60% (diatom-fed), 68% (A. falcatus-fed), to 70% (fasting treatment), but was 0% in the M. aeruginosa diet treatment. Nucleic acid ratios (RNA:DNA), commonly used as proxies for somatic growth potential, were highest in Diporeia feeding on diatoms and lowest in fasting animals. We conclude that overall condition of Diporeia improved with dietary access to EPA and DHA, but survival was not related to this food quantity and/or quality.     

An integrated assessment of polycyclic aromatic hydrocarbons (PAHs) and benthic macroinvertebrate communities in Isle Royale National Park

Polycyclic aromatic hydrocarbons (PAHs) are a class of persistent organic pollutants that are known carcinogens and mutagens. This research used the sediment quality triad, an integrated weight-of-evidence approach, to evaluate sediment PAH concentrations, sediment toxicity, and benthic community structure at marina and reference sites in Isle Royale National Park, USA. The highest PAH concentrations were measured at marina locations and exceeded threshold effect concentrations (161 μg PAH/g TOC) at one site. Marina locations were dominated by pyrogenic PAHs, indicating anthropogenic sources of these compounds. Survival of the amphipod Hyalella azteca was significantly reduced (p = 0.0320) when exposed to sediments from marinas. Although macroinvertebrate abundance and species richness were similar at marina and reference sites, results of multivariate analyses showed that composition of benthic communities varied among sites. In particular, abundance of the PAH-sensitive amphipod, Diporeia spp. was significantly lower at marina sites compared to reference sites. In contrast to patterns observed for organochlorines (e.g., PCBS, dioxins), biota-sediment accumulation factors for PAHs measured in the burrowing mayfly Hexagenia limbata decreased with increasing K_ow values, suggesting that the more lipophilic compounds were being metabolized. Increased PAH concentrations, shifts in community composition, low survival of H. azteca, and reduced abundances of Diporeia spp. at marina sites were consistent with the hypothesis that PAHs impacted these areas; however, across all sites these effects were relatively subtle. These results emphasize the need to use a weight-of-evidence approach when investigating effects of environmental contaminants that occur at relatively low concentrations.     

Genetic assessment of straying rates of wild and hatchery reared lake sturgeon (Acipenser fulvescens) in Lake Superior tributaries

Natal philopatry in lake sturgeon (Acipenser fulvescens) has been hypothesized to be an important factor that has lead to genetically distinct Great Lakes populations. Due to declining abundance, population extirpation, and restricted distribution, hatchery supplementation is being used to augment natural recruitment and to reestablish populations. If hatchery-reared lake sturgeon are more likely to stray than naturally produced individuals, as documented in other well-studied species, outbreeding could potentially jeopardize beneficial site-specific phenotypic and genotypic adaptations. From 1983 to 1994, lake sturgeon propagated using eggs taken from Lake Winnebago adults (Lake Michigan basin) were released in the St. Louis River estuary in western Lake Superior. Our objective was to determine whether these introduced individuals have strayed into annual spawning runs in the Sturgeon River, Michigan. Additionally, we estimated a natural migration rate between the Sturgeon River and Bad River, Wisconsin populations. Presumed primiparous lake sturgeon sampled during Sturgeon River spawning runs from 2003 to 2008 were genotyped at 12 microsatellite loci. Genotypic baselines established for the Sturgeon River (n = 101), Bad River (n = 40), and Lake Winnebago river system (n = 73) revealed a relatively high level of genetic divergence among populations (mean F_ST = 0.103; mean R_ST = 0.124). Likelihood-based assignment tests indicated no straying of stocked Lake Winnebago strain lake sturgeon from the St. Louis River into the Sturgeon River spawning population. One presumed primiparous Sturgeon River individual likely originated from the Bad River population. Four first-generation migrants were detected in the Sturgeon River baseline, indicating an estimated 3.5% natural migration rate for the system.     

Lyngbya wollei in western Lake Erie

We report on the emergence of the potentially toxic filamentous cyanobacterium, Lyngbya wollei as a nuisance species in western Lake Erie. The first indication of heavy L. wollei growth along the lake bottom occurred in September 2006, when a storm deposited large mats of L. wollei in coves along the south shore of Maumee Bay. These mats remained intact over winter and new growth was observed along the margins in April 2007. Mats ranged in thickness from 0.2 to 1.2 m and we estimated that one 100-m stretch of shoreline along the southern shore of Maumee Bay was covered with approximately 200 metric tons of L. wollei. Nearshore surveys conducted in July 2008 revealed greatest benthic L. wollei biomass (591 g/m² ± 361 g/m² fresh weight) in Maumee Bay at depth contours between 1.5 and 3.5 m corresponding to benthic irradiance of approximately 4.0–0.05% of surface irradiance and sand/crushed dreissenid mussel shell-type substrate. A shoreline survey indicated a generally decreasing prevalence of shoreline L. wollei mats with distance from Maumee Bay. Surveys of nearshore benthic areas outside of Maumee Bay revealed substantial L. wollei beds north along the Michigan shoreline, but very little L wollei growth to the east along the Ohio shoreline.     

A short-term look at potential changes in Lake Michigan slimy sculpin diets

Diporeia hoyi and Mysis relicta are the most important prey items of slimy sculpins (Cottus cognatus) in the Great Lakes. Slimy sculpins were collected from dreissenid-infested bottoms off seven Lake Michigan ports at depths of 27–73 m in fall 2003 to study their lake-wide diets. Relatively large dreissenid biomass occurred at depths of 37- and 46-m. Quagga mussels (Dreissena bugnesis) composed at least 50% of dreissenid biomass at Manistique, Saugatuck, and Sturgeon Bay. Mysis accounted for 82% of the sculpin diet by dry weight at eastern Lake Michigan while Diporeia composed 54–69% of the diet at western Lake Michigan and dominated the diets of slimy sculpins at all sites deeper than 46 m. In northern Lake Michigan, this diet study in new sites showed that slimy sculpin consumed more prey with low energy contents, especially chironomids, than Mysis and Diporeia in shallow sites (depth <55 m). We recommend diet studies on sedentary benthic fishes to be conducted along perimeters of the Great Lakes to observe changes in their diets that may be impacted by changing benthic macroinvertebrate communities.     

Changes in Fecundity and Egg Lipid Content of Lake Whitefish (Coregonus clupeaformis) in the Upper Laurentian Great Lakes between 1986–87 and 2003–05

Lake whitefish (Coregonus clupeaformis Mitchill), an important commercial species in the Laurentian Great Lakes, have experienced decreased growth and condition in regions of the upper Great Lakes over the past 20 years. Increases in lake whitefish density and decreases in the density of Diporeia spp., an energy rich and historically important part of the lake whitefish diet, have been implicated in the recent declines in lake whitefish growth and condition. The goal of this study was to describe lake whitefish fecundity, egg lipid content, and total ovary lipid content in selected regions of Lakes Huron, Michigan, and Superior in 1986–87 and 2003–05, two time periods with different lake whitefish and Diporeia densities. Under conditions of high lake whitefish density and low Diporeia density, female lake whitefish in the upper Laurentian Great Lakes generally produced fewer eggs. Egg lipid content was higher in 2003–05 than in 1986–87 at all sites, regardless of changes in lake whitefish or Diporeia densities. Total ovary lipid content and lake whitefish abundance were inversely related, while there was no significant relationship between total ovary lipid content and Diporeia density. The amount of energy that lake whitefish invested in egg production was more closely associated with lake whitefish abundance than with Diporeia density. This study provides evidence that recent changes in production dynamics of Great Lakes lake whitefish have not been driven solely by declines in Diporeia but have been significantly influenced by lake whitefish abundance.     

Stock assessment and fishery management of Henicorhynchus spp., Cyclocheilichthys enoplos and Channa micropeltes in Tonle Sap Great Lake,Cambodia

Tonle Sap Great Lake, in the lower Mekong River basin, contributes 60% of Cambodia's inland fisheries catch. There are four types of fisheries, including the middle-scale fishery. The major species comprising the middle-scale fishery are Henicorhynchus spp. (16%), a small-bodied fish caught mainly with small-mesh nets, and Cyclocheilichthys enoplos (13%), and Channa micropeltes (7%) which are caught with a variety of gear. Here we apply the Schaefer surplus production model to the middle-scale fisheries using monitoring data collected between 1995 and 1999. Model simulations determined that the maximum sustainable yield (F_MSY) for Henicorhynchus spp., was obtained with the use of 47,206 gillnets (mesh size < 50 mm) and 4269 and 1605 fishing boats for C. enoplos and C. microplestes, respectively. Over 1995–1999, there was a strong trend of increase in fishing effort and decrease of catch-per-unit-effort in the waters of Pursat, Siem Riep and Kampong Chnnang provinces. These provinces have large population centres located close to the fishing grounds; fishing in these areas should be more strictly regulated.