First documentation of spawning and nest guarding in the laboratory by the invasive fish,the round goby (Neogobius melanostomus)

Although the round goby, Neogobius melanostomus, is widespread in the Great Lakes and has an extended breeding season with a high reproductive rate; its spawning behaviour remains elusive. We present the first reported accounts of spawning by the round goby in the laboratory. By simulating winter conditions and restoring spring conditions, we induced round gobies to spawn in October 2007, March 2008, May 2008, and January 2009. In one case, fanning by the nest-holding male began 10 days before egg deposition and, during this period, the male rubbed secretions along the ceiling of the nest. Males were choosy about which gravid females entered the nest and prevented entry by some females. Spawning involved repeated inversions by females and males releasing gametes on the ceiling of the nest. Males guarded the nest by blocking the entrance, producing agonistic vocalizations and chasing intruders. Inside the nest, eggs were regularly inspected by the males and constantly ventilated using pectoral and caudal fins. Up to three gravid females spawned sequentially in a nest. Peak ventilation occurred after egg deposition and declined with time until the parental male ate the eggs. The decline of parental care and egg cannibalism was likely an artifact of laboratory conditions and small brood size. Our findings offer new information on the reproductive habits of the invasive round goby. Because the reproductive sequence in the laboratory seems easy to disrupt, the procedures may lead to a management tool to control the spread of the species into new areas.     

Response to putative round goby (Neogobius melanostomus) pheromones by centrarchid and percid fish species in the Laurentian Great Lakes

Pheromone trapping is an increasingly viable strategy to reduce invasive fish populations, largely due to the pheromones' function of evoking behavioral responses among conspecifics. Prior to attempting such population control techniques, the pheromones must be identified and their possible influences on non-target species addressed. The round goby (Neogobius melanostomus) is a species invasive to the Great Lakes region, and negatively impacts the ecosystem by interfering with local fish populations. At least two 5β-reduced 3α-hydroxyl steroids released by reproductive N. melanostomus (11-O-ETIO and 11-O-ETIO-3s) evoke olfactory sensory responses from the olfactory epithelium of conspecifics, and water conditioned by reproductive males (containing these steroids) attracts female round gobies. In this study, we examined whether these putative pheromones, along with simultaneously-released 11-O-ETIO-17s, stimulate olfactory sensory responses from alternative fish species sharing the same ecosystem as N. melanostomus in the Great Lakes region. Rock bass (Ambloplites rupestris), bluegill sunfish (Lepomis macrochirus), pumpkinseed sunfish (Lepomis gibbosus), smallmouth bass (Micropterus dolomieu), and yellow perch (Perca flavescens) were the targets of an electro-olfactogram experiment designed to record responses to odors. When compared to round goby responses from previous studies, amino acids and the bile acid consistently elicited electro-olfactogram responses across all species, but only round gobies showed a response to the putative pheromones. This study supports the concept of conducting a pheromone trapping trial in the field without adversely affecting the olfactory responses of non-target fish in the area.     

Size-selective predation by round gobies (Neogobius melanostomus) on dreissenid mussels in the field

Although numerous studies have shown that round gobies (Neogobius melanostomus) prey on dreissenid mussels (Dreissena polymorpha and Dreissena bugensis), there is an apparent shortage of detailed field studies on the subject. The 5-month field study described here quantifies predation by round gobies on dreissenids in Presque Isle Bay, Lake Erie. Dreissenids dominated the diet of round gobies, composing 92% of the prey items recovered. Over half of the 3870 valves (1935 mussels) recovered from 155 round gobies were crushed, while the remainder were swallowed whole. Crushed dreissenids were larger than those swallowed whole, and the tendency to crush dreissenids did not vary among three length classes of round goby. Round goby length was positively related to average size of dreissenids consumed, average size of whole and crushed dreissenids, largest whole dreissenid consumed, and largest crushed dreissenid consumed. Indices of selectivity revealed similarly shaped curves for three length classes of round gobies, a shift toward larger dreissenid size classes with an increase in round goby length, and peak preferences for 8-11 mm dreissenids. Factors such as gape limitation, availability and accessibility of differently sized dreissenids, forces generated while removing mussels from the substrate and crushing them, and caloric content of dreissenids all likely play roles in the observed size-selectivity and differential processing of dreissenids. Although factors influencing size-selectivity are not completely understood, the observed preference of round gobies for dreissenids near the size when they are first reproductive could impact the demography of dreissenid populations.     

Effectiveness of Piscicides for Controlling Round Gobies (Neogobius melanostomus)

Round gobies (Neogobius melanostomus) were introduced to the Great Lakes presumably as a result of ballast water releases from seagoing freighters returning from European water bodies. These unwelcome fish have become established in the Great Lakes region and are expanding their range to suitable portions of other interior drainage basins including the Mississippi River traversing the central United States and the Trent-Severn waterway spanning south-central Ontario. If the invasion continues, use of chemical toxicants as a control measure may be necessary. Toxicity tests of the currently registered piscicides antimycin, rotenone, 3-trifluoromethyl-4-nitrophenol (TFM), and Bayluscide^® were conducted with three fish species native to the Great Lakes and round gobies collected from the Illinois Waterway. Tests indicated that round gobies are sensitive to all of the piscicides, however, the level of sensitivity is similar to that of the native fish species tested. Therefore, currently registered piscicides have limited potential to selectively remove round gobies. Bottom-release formulations of Bayluscide^® and antimycin were also evaluated as control agents for the normally bottom-dwelling round goby. Avoidance behavior tests demonstrated that the round goby did not react to the presence of either chemical. Therefore, the bottom-release formulations may have some application for the selective removal of round gobies, and may be one of the few tools presently available to fishery managers to help limit the range expansion of this invasive fish.     

Density and habitat use by the round goby (Apollonia melanostoma) in the Bay of Quinte,Lake Ontario

We assessed round goby (Apollonia melanostoma) density and size structure in two sections of the Bay of Quinte (Lake Ontario) that had been invaded by this species two years apart. Round goby density was assessed with 50 m linear transects, recorded with an underwater video recording apparatus developed for this study that included a depth sounder for maintaining a fixed distance above the substrate. The highest mean round goby densities were observed in the shallowest depth zone (1.5–3 m) at both sites, but there were differences between the sites in the habitat types where the highest densities occurred and there were no significant density differences among habitat types at either site (rock with sparse vegetation, mud with sparse vegetation, sand/mud with moderate vegetation cover). In the upper bay, mean body length of round gobies declined with depth, whereas in the lower bay, mean round goby length was greatest in the deepest zone. Mean body length of round gobies did not differ significantly by habitat type in either section of the bay.     

Double-crested Cormorant (Phalacrocorax auritus) chick bioenergetics following round goby (Neogobius melanostomus) invasion and implementation of cormorant population control

As the fish community changed in the Beaver Archipelago (northern Lake Michigan), so has the diet of breeding Double-crested Cormorants (Phalacrocorax auritus). In 2000, the energetically dense alewife (Alosa pseudoharengus) dominated the diet, but more recently, the relatively low quality round goby (Neogobius melanostomus) has become the most common prey item. Additionally, cormorant control measures have been underway in the archipelago. This study investigated decreases in cormorant chick numbers and changes in chick bioenergetics in response to this change in diet, as well as the influence of control efforts, and compared these results to an earlier study. Two colonies, one actively controlled the other only controlled following completion of this study, were investigated to determine changes in colony size throughout the breeding season, document chick diet, measure chick growth, and determine chick survivorship to fledging. The bioenergetics model estimated that in 2010, chicks consumed a greater biomass of prey to reach fledging due to the change in diet, but lower biomass overall due to fewer chicks in the system. Control efforts in combination with the change in diet reduced chick numbers. Overall, the impact of cormorant chicks on available fish biomass has declined in recent years due to control efforts. However, the birds have the potential to consume a greater biomass of round goby in the absence of control. The impact of cormorants on fisheries resources is complicated by the abundance and effects of non-native species.     

Round goby (Neogobius melanostomus) population structure,biomass, prey consumption and mortality from predation in the Bay of Quinte,Lake Ontario

We compared round goby (Neogobius melanostomus) population structure and biomass, estimated the quantity of prey consumed by round goby and assessed predation on round goby in two areas of the Bay of Quinte (upper and lower bay) that were invaded by this species two years apart. A bioenergetics approach was used to estimate round goby consumption and to illustrate the influence of changes in the round goby biomass, from May to October in both areas of the bay. Mean round goby biomass was 5 tonnes/km² for the upper bay and 11.2 tonnes/km² for the lower bay. Round goby populations were dominated by age-0 to age-2 individuals; older individuals were rare although they constituted a greater portion of total biomass in the earlier invaded lower bay than the upper bay. Estimated round goby consumption was 78.4 tonnes/km² for the upper bay and 127.1 tonnes/km² for the lower bay; dreissenids were the most important prey followed by chironomids, zooplankton and amphipods. Diet analysis showed that round gobies were consumed by all predator species analyzed, and constituted a high percentage of stomach content biomass in yellow perch (Perca flavescens), largemouth bass (Micropterus salmoides) and northern pike (Esox lucius). Age structure differences between the upper and lower bay are consistent with those expected between expanding and stable populations.     

Multiple male reproductive morphs in the invasive round goby (Apollonia melanostoma)

Alternative male reproductive tactics are taxonomically widespread. In such species, parental, or conventional, males express secondary sexual characteristics, court females and guard offspring, while smaller parasitic or sneaker males avoid the costs of courtship and parental care by performing sneak fertilizations. Theory predicts that sneakers will invest more in testes mass and produce more competitive ejaculates than parentals because sneakers always experience sperm competition while parental males experience sperm competition only when a sneaker is present. Here we present convergent lines of evidence supporting the existence of alternative male reproductive tactics in round gobies (Apollonia melanostoma, formerly Neogobius melanostomus), a recent invader in the Great Lakes. Dark morph males exhibited secondary sexual characteristics, were larger and had higher plasma 11-ketotestosterone concentrations than light morphs, while light morph males invested more in ejaculates (both testes mass and sperm density). Both male morphs had enlarged urogenital papillae, but papillae were relatively longer in light morph males. Sperm tail length did not differ between morphs, and sperm from dark morphs swam faster than sperm from light morphs. Our data strongly argue for the presence of alternative tactics in round gobies, support some predictions from sperm competition theory and align with empirical observations in other taxa. For species of concern like the invasive round goby, it is critical to consider such evidence of alternative male mating tactics when constructing population growth models and assessment of invasion success and impacts.     

Ontogenetic changes in pharyngeal morphology correlate with a diet shift from arthropods to dreissenid mussels in round gobies (Neogobius melanostomus)

The potential of predators to regulate populations of dreissenid mussels (Dreissena polymorpha and Dreissena rostriformis bugensis) has been addressed since early in the dreissenid invasion of North America. Round gobies (Neogobius melanostomus) larger than approximately 60 mm have been shown to prey extensively on dreissenids, whereas smaller round gobies feed mainly on aquatic insects and crustaceans. We propose that ontogenetic changes in pharyngeal morphology may contribute to this diet shift in round gobies. Pharyngeals of 69 round gobies ranging from 31 to 164 mm total length were investigated using light microscopy and scanning electron microscopy. Areas of lower pharyngeals and pharyngobranchial 2 increased allometrically with fish length. Pharyngeals of round gobies smaller than 50 mm contained narrow (< 0.1 mm diameter) papilliform teeth that are consistent with eating soft-bodied prey. By the time round gobies reached approximately 80 mm in length, pharyngeals contained larger diameter (0.3-0.5 mm) molariform teeth typical of those found in molluscivorous fish. Pharyngeal teeth of the largest round gobies also showed considerable wear. Although changes in pharyngeal morphology may contribute to the previously described diet shift in round gobies as they age, genetic and environmental factors both likely influence pharyngeal remodeling and therefore merit further investigation.     

Evaluating gonadosomatic index as an estimator of reproductive condition in the invasive round goby, Neogobius melanostomus

Using gonadosomatic index cut-off scores has become a standard protocol for selecting reproductive fish in studies on the reproductive biology of round goby, Neogobius melanostomus, a significant invader of the Laurentian Great Lakes, but the validity of this practice has not been validated with histological staging. The goal of the current study was to evaluate the effectiveness of using gonadosomatic index (GSI) cut-off scores to classify reproductive status in male and female round goby by documenting associations between GSI, sex steroids, and gonad development. Gonadal stage was determined in both sexes using hematoxylin and eosin histology. Plasma 11-ketotestosterone and testosterone were measured in males, and testosterone and 17β-estradiol were measured in females. Gonadosomatic index cut-off scores were effective in selecting spawning capable individuals at higher GSI values, but GSI values were limited in the ability to make further distinctions of gonadal stage and missed many spawning capable females. In females, testosterone levels were highest during vitellogenic growth and declined prior to ovulation. 17β-estradiol displayed a similar, but non-statistically significant pattern. Males with developing testes had higher levels of 11-ketotestosterone —but not testosterone —than reproductively immature males, although levels of these androgens were overall positively correlated in males. The findings indicate that the conventional GSI cut-off scores (1% in males, 8% in females) accurately assign spawning capable condition in both sexes; however, they may also exclude some spawning capable females with lower GSIs.     

Impacts of predation by the Eurasian round goby (Neogobius melanostomus) on molluscs in the upper St. Lawrence River

An invasive Eurasian fish, the round goby Neogobius melanostomus, has recently spread from the Great Lakes into the St. Lawrence River. We quantified prey preferences of this benthivore and determined whether its predatory impacts on molluscs in the river are similar to those in the Great Lakes. We measured the size structure of gastropods and dreissenid mussels at 13 St. Lawrence River sites where round goby densities ranged from 0 to 6 m^− 2. For four of these sites, data were available for multiple years before and after invasion. Contrary to studies in the Great Lakes, there were no consistent effects of round goby density on the size structure of dreissenids, although there was an ontogenetic diet shift toward dreissenids. However, the abundance and richness of small gastropods (≤ 14 mm) was negatively correlated with round goby density across all sites, and declined over time at three of four sites sampled before and after invasion. Median gastropod size also declined across sites with increasing round goby density. Gastropods (as well as chironomid larvae, caddisfly larvae, and ostracods) were consistently among the most preferred prey items consumed by gobies, whereas dreissenids (as well as leeches and freshwater mites) were consistently avoided. These results indicate the major role of the round goby in structuring gastropod populations in the St. Lawrence River, and highlight large-scale spatial variation in its predatory impact on dreissenid populations.     

First report of tubenose goby (Proterorhinus semilunaris) in the eastern basin of Lake Erie

The tubenose goby (Proterorhinus semilunaris), native to the Ponto-Caspian region, was first discovered in the Laurentian Great Lakes in 1990 after it was introduced through ballast water discharge. Compared with Neogobius melanostomus, another exotic gobiid from the Ponto-Caspian, colonization of the Great Lakes by P. semilunaris has been slow, with reports of the species being largely confined to the Huron-Erie Corridor (HEC) and western portions of Lake Erie and Lake Superior. This is the first report of P. semilunaris in the Great Lakes east of the western basin of Lake Erie. Between 28 June and 27 July, 2012, 176 P. semilunaris were collected from shallow (< 1.2 m) water of Marina Lake, a 40 ha embayment in Presque Isle State Park (Erie, PA). The large number of P. semilunaris collected at the site and the presence of individuals as small as 17 mm total length suggest an established population. However, the mechanism by which P. semilunaris was introduced to Presque Isle Bay is not clear.     

Laboratory evaluation of spawning substrate type on potential egg predation by round goby (Neogobius melanostomus)

Spawning habitat structure may protect demersal eggs of broadcast spawning species from depredation. Egg predation by round goby (Neogobius melanostomus) is commonly referenced as a concern associated with their invasion of the Laurentian Great Lakes. Whereas nest-building species have received some attention in egg predation studies, broadcast spawning species may be particularly vulnerable as they do not guard their eggs. This study used a 2?×?7 factorial experiment to investigate how substrate characteristics influenced the rate that fertilized eggs are lost when exposed to round goby. Eggs for two recreationally important broadcast spawning species, northern pike (Esox lucius) and muskellunge (Esox masquinongy), were placed within spawning habitat treatments (bare, silt, sand, rubble, gravel, filamentous algae, and submerged aquatic vegetation) representing a range (least to most complex) of habitat complexity. Regardless of substrate type, egg loss was similar between the two esocid species. Across all substrates and species, the number of eggs lost varied two-fold over the 24-hour experiment, with the lowest rate of egg loss observed in the most complex substrate (submerged aquatic vegetation) and the highest over the least complex substrates (bare and silt). Both northern pike and muskellunge are known to spawn over structurally complex submerged aquatic vegetation and filamentous algae. Although eggs spawned over these substrates likely offer some protection from predation, eggs that settle over less complex ancillary habitats may face higher predation risk.     

First records of ratan goby Ponticola ratan in the Kaniv Reservoir and Desna River (Ukraine): A discreet invader of rip-raps

Monitoring of nearshore fish communities carried out in 2019 and 2022 in the Kaniv Reservoir and lower Desna River (Dnieper River basin, Ukraine) showed the presence of the ratan goby (Ponticola ratan), which has never been observed in these water bodies before and naturally inhabited marine and brackish waters of the Black and Azov Sea basins. The ratan goby was detected only on rip-rap structures and not on any other surveyed substrates including sand, mud, clay, and gravel which suggests a narrow habitat preference for this species. In all instances, the ratan goby co-occurred with the round goby (Neogobius melanostomus). Occurrence on all surveyed rip-raps, high abundances and the presence of a variety of size classes of the ratan goby in samples suggest that this species penetrated the Kaniv Reservoir quite long ago and successfully established there and has become a common component of the local fish fauna. Based on the obtained data, this gobiid should be listed among freshwater fishes of Europe as a euryhaline species that can potentially colonize other rivers flowing into the Black Sea including the Danube.     

Fish and Macroinvertebrate Communities in Tributary Streams of Eastern Lake Erie with and without Round Gobies (Neogobius melanostomus, Pallas 1814)

Round gobies have had significant impacts on benthic fish and invertebrate communities in nearshore habitats of the Great Lakes. As round gobies have become more abundant in lake habitats, there has been an expansion of their populations into tributary streams and rivers. We compared stream invertebrate and fish communities in New York tributaries to Lake Erie with round gobies present and absent. Four of six benthic invertebrate metrics differed between streams with and without round gobies. Streams with round gobies present had reduced Shannon diversity, EPT richness, and EPT/chironomid ratios, and increased macroinvertebrate density relative to streams without round gobies, but there was no difference in non-Diptera density, or total taxa richness. None of the four fish metrics examined differed between streams with and without round gobies. However, darters occurred in all streams lacking round gobies, but did not occur in any streams with round gobies. Comparisons with historical fish and macroinvertebrate distributional data support our suspicion of goby-induced community changes. In these New York streams, round gobies seem to have had significant impacts on invertebrate communities via their consumptive behavior, whereas the impacts on fish communities are less evident. If round gobies continue to expand their distribution inland, the resultant alterations in macroinvertebrate communities may impact the suitability of tributary streams as spawning and nursery habitat for several sport fish species and for energy dynamics in tributary streams.     

Slimy sculpin depth shifts and habitat squeeze following the round goby invasion in the Laurentian Great Lakes

The collapse of Diporeia spp. and invasions of dreissenid mussels (zebra, Dreissena polymorpha; quagga, D. bugensis) and round goby (Neogobius melanostomus) have been associated with declines in abundance of native benthic fishes in the Great Lakes, including historically abundant slimy sculpin (Cottus cognatus). We hypothesized that as round goby colonized deeper habitat, slimy sculpin avoided habitat competition, predation, and aggression from round goby by shifting to deeper habitat. Accordingly, we predicted increased depth overlap of slimy sculpin with both round goby and deepwater sculpin (Myoxocephalus thompsonii) that resulted in habitat squeeze by both species. We used long-term bottom trawl data from Lakes Michigan, Huron, and Ontario to evaluate shifts in slimy sculpin depth and their depth overlap with round goby and deepwater sculpin. Lake Huron most supported our hypotheses as slimy sculpin shifted to deeper habitat coincident with the round goby invasion, and depth overlap between slimy sculpin and both species recently increased. Slimy sculpin depth trends in Lakes Michigan and Ontario suggest other ecological and environmental factors better predicted sculpin depth in these lakes.     

A lakewide survey of Asian clam (Corbicula fluminea) distribution at warmwater discharges in Lake Michigan

The Asian clam Corbicula fluminea is among the most prolific aquatic invaders in the world; but in colder mid-latitude areas, like the Laurentian Great Lakes, their population expansion has likely been limited by poor overwinter survival. In these areas, Asian clams are typically found in thermal refugia like warmwater discharges from industrial facilities. We sought to identify the current extent of Asian clam populations in Lake Michigan and waters immediately adjacent to it, specifically at locations most likely to harbor overwintering populations – industrial warmwater discharges. During April–May 2017, we surveyed 17 locations around Lake Michigan. Evidence of Asian clam populations was found at four sites, though live specimens (n?=?3) were only found at the Indiana Harbor Ship Canal in East Chicago, IN. Shells or fragments of shells were found at Green Bay, WI, Waukegan, IL, and Port Sheldon, MI. Our findings indicate that although Asian clams are present in Lake Michigan, they are relatively rare, and remain isolated to a few small pockets of over-wintering habitat.     

Factors governing the distribution and fish-community associations of the round goby in Michigan tributaries of the Laurentian Great Lakes

The round goby (Neogobius melanostomus) is increasingly being reported in tributaries of the Laurentian Great Lakes where these fish have been shown to adversely impact native stream biota. Determining the characteristics and distribution of invaded streams are the first steps toward effective round goby management. We sampled 30 tributaries in the Great Lakes basin and characterized each in terms of nine physical reach-scale attributes. Round goby were detected in 14 streams where abundances ranged from 4% to 53% of the fish sampled in each stream. Round goby was the single most abundant fish species sampled, constituting 14% of all fish encountered across all sites, and 30% of individuals in round goby-present sites. Round goby-present sites were larger, had lower channel slopes, less large wood, and less canopy cover than round goby-absent sites, suggesting that these attributes may promote round goby establishment. Mottled sculpin, cyprinids, brook stickleback, white sucker and rainbow trout were associated with goby absence while centrarchids, percids, yellow bullhead, and mud minnow were associated round goby presence. Collectively these results demonstrate that round goby are widespread in eastern Michigan tributaries to the Great Lakes, present in streams with a range of physical habitat characteristics, and that round goby presence is associated with certain fish species.     

Quantifying a shift in benthic dominance from zebra (Dreissena polymorpha) to quagga (Dreissena rostriformis bugensis) mussels in a large,inland lake

Dreissenid mussels are aggressive invasive species that are continuing to spread across North America and co-occur in the same waterbodies with increasing frequency, yet the outcome and implications of this competition are poorly resolved. In 2009 and 2015, detailed (700 + sample sites) surveys were undertaken to assess the impacts of invasive dreissenid mussels in Lake Simcoe (Ontario, Canada). In 2009, zebra mussels were dominant, accounting for 84.3% of invasive mussel biomass recorded. In 2015, quagga mussels dominated (88.5% of invasive mussel biomass) and had expanded into profundal (> 20 m water depth) sites and onto soft (mud/silt) substrates with a mean profundal density of 887 mussels/m² (2015) compared to ~ 39 mussels/m² in 2009. Based on our annual benthos monitoring, at a subset of ~ 30 sites, this shift from zebra to quagga mussels occurred ~ 2010 and is likely related to a population decline of zebra mussels in waterbodies where both species are present, as recorded elsewhere in the Great Lakes Region. As the initial invasion of dreissenid mussels caused widespread ecological changes in Lake Simcoe, we are currently investigating the effects this change in species dominance, and their expansion into the profundal zone, will have on the lake; and our environmental management strategies. Areas of future study will include: changes in the composition of benthos, fish, or phytoplankton communities; increased water clarity and reduction of the spring phytoplankton bloom; energy/nutrient cycling; and fouling of anthropogenic in-lake infrastructures (e.g. water treatment intakes) built at depths > 25 m to avoid previous zebra mussel colonization.     

Application of a numerical model for circulation,temperature and pollutant distribution in Hamilton Harbour

The restoration of Hamilton Harbour, from an environmental standpoint, is a current concern for the agencies involved with remediation efforts in the harbour. Estimates of circulation and mixing are needed to assess the fate and transport of water quality constituents in the harbour. A three-dimensional hydrodynamic modeling system (ELCOM) is used to study the circulation and thermal structure in the harbour. The model results were compared with profiles of temperature at several moorings and currents and water levels in the harbour. The model showed considerable skill in reproducing the thermal structure, surface currents and water levels. Mean summer circulation in the harbour showed two counter-rotating gyres occupying the harbour. The model produced harbour-lake exchange characteristics are in agreement with previous studies. Simulations using passive tracers qualitatively agreed with chemical tracer studies conducted near a sewage treatment plant outfall. The accuracy of these simulations suggests that the model is capable of describing flow and transport of material required for detailed water quality simulations.