Menidia audens

Invasion History

First Non-native North American Tidal Record: 1971
First Non-native West Coast Tidal Record: 1971
First Non-native East/Gulf Coast Tidal Record:

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General Invasion History:

Mississippi Silverside (Menidia audens) is a freshwater fish that is closely related to Inland Silverside (M. beryllina), an estuarine fish which enters tidal freshwater, and the lower reaches of nontidal rivers. The Inland Silverside ranges from Cape Cod, Massachusetts to northeastern Mexico (Robins et al. 1986). Mississippi Silversides occur in the Mississippi and Gulf basins, from Louisiana-Texas to southern Illinois, westward to Arkansas and Oklahoma (Page and Burr 1991; Suttkus and Thompson 2002; Suttkus 2005; Fluker et al. 2011). The Mississippi Silverside has been introduced to reservoirs on the Tennessee and Cumberland Rivers in Alabama, Tennessee, and Kentucky, as well as to many lakes and reservoirs in California (Dill and Cordone 1997; USGS Nonindigenous Aquatic Species Program 2018). Mississippi Silversides were introduction to the Upper and Lower Blue Lake and Clear Lake in California in 1967 and 1969, the San Joaquin River in 1971. They spread rapidly through the San Francisco Bay estuary becoming one of its most abundant species. Through the California Aqueduct System, they were able to colonize reservoirs and river systems in Southern California, eventually reaching estuaries and lagoons (Swift 2014). The Mississippi Silverside has retained tolerance to marine salinities, and occurs in San Francisco Bay salt ponds with salinities of 25 to 30 PSU (Mejia et al. 2008), and marine embayments in Southern California (Swift 2014). Beginning in 1990, Mississippi Silversides colonized reservoirs in the Tennessee and Cumberland River systems, east of their native range, through the Tennessee-Tombigbee Waterway barge-canal (Suttkus et al. 2005; USGS Nonindigenous Aquatic Species Program 2018).

North American Invasion History:

Invasion History on the West Coast:

In 1967, 9,000 Mississippi Silversides from Lake Texoma, Oklahoma were introduced to Clear Lake and Lower and Upper Blue Lakes in central California for control of the Clear Lake Gnat (Chaoborus astictopus) and algae. The introductions into the Blue Lakes in the Sierra were made with permission from the California Fish and Game Commission, but the stocking of Clear Lake was reportedly an afterthought and not authorized by the commission. Clear Lake's outflows into Cache Creek, which is a tributary of the Sacramento River and provides a route to the San Francisco Estuary Delta. Mississippi Silversides were found in Cache Creek and Putah Creek in 1972 and were reproducing in the Delta by 1975. By 1980, the silversides had become one of the dominant species in the Sacramento-San Joaquin Delta (Cohen and Carlton 1995; Dill and Cordone 1997; Moyle 2002). While Mississippi Silversides are associated with the low-salinity regions of the Delta, they also occur in Napa River saltmarshes in salinities of 17-18 PSU (Cohen et al. 2008) and in South Bay salt ponds, which have mean salinities of 25-50PSU (Mejia et al. 2008). In a 2014 survey of the Sacramento-San Joaquin Delta, Mississippi Shad and Threadfin Shad made up 43% of the nekton abundance (Feyrer et al. 2017). In freshwater portions of the Delta, Mississippi Silversides showed a steady pattern of increase from 1996 to 2011, possibly levelling off after 2011 (Mahardja et al. 201).

From 1988 to 1992 Mississippi Silversides were caught in Silverwood, Castaic, Pyramids Lakes, and other reservoirs in southern California. This fish were probably transported from the Delta by the California Aqueduct (Swift et al. 1993). From 1997 to 2001 larval and juvenile Mississippi Silversides were caught in King Harbor, Redondo Beach, and Los Angeles County. No further catches were made there, but subsequently, this fish was found in coastal lagoons in Santa Barbara (2002) and in the San Gabriel River (2006-2007). Populations in lagoons at the mouth of the Santa Clara River (2006-2012) and Malibu Lagoon (2005-2013) appear to be established. Most of these introductions appear to have come from fish transported to southern California streams by the California Aqueduct, where untreated water enters local streams, but in other streams introduction as discarded bait is likely (Swift et al. 2014).

Invasion History on the East Coast:

In 1990, Mississippi Silverside (Menidia audens) was collected in Barkley Reservoir, on the Cumberland River, Tennessee, east of its native range. It was collected in 1991 in the Kentucky Reservoir, on the Cumberland River, Kentucky, and in 1993, in the Pickwick Reservoir, on the Tennessee River, Alabama (Suttkus et al. 2005; Simmons 2013; USGS Nonindigenous Aquatic Species Program 2018). These reservoirs are connected by the Tennessee-Tombigbee Waterway, a barge canal system. Fish probably entered the reservoirs using the Ohio and Tennessee River, using navigation locks (Simmons 2013).

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Description

The Mississippi Silverside, Menidia audens (Hay 1882) is a small freshwater fish, belonging to a species complex that includes the Inland Silverside (Cope 1867, Menidia beryllina) and the Tidewater Silverside (Goode & Bean, 1879, Menidia peninsulae). The Island Silverside is estuarine, ranging from marine salinities to freshwater, while the Tidewater Silverside is more marine (Robins et al. 1986; Fluker et al. 2011). The Mississippi Silverside ranges in the lower Mississippi basin from southern Illinois, and westward to Arkansas and Oklahoma, to tidal brackish waters of the Mississippi, where it overlaps with the Inland Silverside (M. beryllina). The populations of Mississippi and Inland Silversides are closely related but genetically and morphologically divergent, with a sharp distinction in phenotype and mtDNA at the freshwater-brackish transition. This is indicative of a recent invasion of freshwater and rapid evolution (Fluker et al. 2011). The taxonomic status of Mississippi Silverside has fluctuated from distinct species to subspecies of the Island Silverside to complete synonymy and back again (Chernoff et al. 1991; Suttkus and Thompson 2002; Sutkuss 2005; Fluker et al. 2011). Fluker et al. (2011) are tentative in regarding the species status of Mississippi Silversides. It may be regarded as an emergent species, in the early process of differentiation, and in the process of colonizing fresh water. The Mississippi Silverside retains its tolerance for seawater (Hubbs and et al. 1971; Mejia et al. 2008; Swift et al. 2014).

Silversides are slim, small, silvery fishes with separate spiny and soft dorsal fins, a forked tail, with a silvery band on the side and a lateral line. The first dorsal fin of a Mississippi Silverside (Menidia audens) is anterior to the origin of the anal fin. The mouth is small, and the snout is long, about equal to eye diameter. The anal fin is sickle-shaped (Page and Burr 1991). The first dorsal has 3-4 spines and the second dorsal has 8-9 rays and 16-19 anal rays (Suttkus and Thompson 2002; Suttkus et al. 2005). The Mississippi Silverside and Inland Silverside (Menidia beryllina) differ in many small morphometric and meristic differences. Overall, Mississippi Silversides are slimmer than Inland Silversides have slimmer bodies and a thinner caudal peduncle (Suttkus et al. 2005; Swift et al. 2015). Mississippi Silversides have more predorsal scales (20-23) and lateral line scales (39-45) than Inland Silversides (14-18) and (36-40) (Suttkus and Thompson 2002). Adults can reach 100mm, but more normal maxima are 70-90 mm (Suttkus and Thompson 2002; Page and Burr 1991; California Fish Website 2018). The fish are yellowish green on the back, and a silvery band on the sides, and a transparent green belly (Page and Burr 1991; Moyle 2002; California Fish Website 2018).

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Taxonomy

Ecology

General:

Mississippi Silversides (Menidia audens, Hay 1882) are small freshwater fish, apparently recently evolved from the estuarine fish, M. beryllina, which has retained the ability to colonize brackish and marine waters (Hubbs et al. 1971; Fluker et al. 2011; Swift 2014). Males and females are not morphologically distinguishable, except by the mass of eggs in a female's ovary. Adults in Lake Texoma, Oklahoma begin spawning at ~ 50 mm and very rarely survive to a second year. Spawning in the Delta takes place in April through September, with more than one spawning per year (Wang 1986). Females produce 44,000-59,000 eggs over the course of a season (Mense 1967). The eggs are scattered on vegetation and hatch in 4 to 30 days, depending on temperature. The larvae have a prominent air-bladder. The yolksac is absorbed when the fish reaches 5 mm in length and are fully formed juveniles at 9 mm (Wang 1986). In Lake Texoma, the first juveniles appeared ~2 months after the start of spawning, the maximum age was ~16 months.

Mississippi Silversides apparently tolerant a wide range of temperature and salinity. For example in Lake Texoma, Oklahoma, they survive temperatures ranging from 7 to 30 C (Work and Gophen 1999), and probably experiences lower temperatures in rivers and reservoirs farther north in Kentucky and Tennessee (Simmons et al. 2013). This fish has become established in freshwater, but retains the ability to survive and possibly reproduce in strongly brackish or marine waters. Embryos from Lake Texoma developed in salinities ~9 PSU, while adults tolerated salinities close to seawater (Hubbs et al. 1971). Mississippi Silversides were collected in the Alviso Salt Ponds, San Francisco Bay, at salinities of ~24 to 50 PSU (Mejia et al. 2008), and have colonized marine embayments and lagoons in southern California (Swift 2014). Reported optimal salinities for Mississippi Silversides range from 3.4 PSU (Mahajarda et al. 2015) to 10-15 PSU (Moyle 2002).

Mississippi Silversides swim in large schools, preferring shallow, open waters over sand and gravel (Moyle 2002). They are carnivores, feeding on zooplankton, insects, small benthic invertebrates, and small fishes, including the endangered Delta Smelt (Hypomesus transpacificus) (Cohen and Bollens 2008). As a small schooling fish, they are also prey for a wide variety of predators, including the introduced Striped Bass (Morone saxatilis) and Largemouth Bass (Micropterus salmoides) and the native Sacramento Pikeminnow (Ptychocheilus grandis), and aquatic birds, such as the Western Grebe (Aechmophorus occidentalis) (Moyle 2002; Nobriga and Feyrer 2007).

Food:

copepods, cumaceans, flying insects, Delta Smelt

Consumers:

Fishes

Competitors:

Threadfin Shad; Wakasagi. Delta Smelt

Trophic Status:

Carnivore

Carn

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Habitats

General Habitat	Fresh (nontidal) Marsh	None
General Habitat	Grass Bed	None
General Habitat	Nontidal Freshwater	None
General Habitat	Tidal Fresh Marsh	None
General Habitat	Unstructured Bottom	None
General Habitat	Salt-brackish marsh	None
Tidal Range	Subtidal	None
Vertical Habitat	Nektonic	None

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Life History

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Tolerances and Life History Parameters

Minimum Temperature (ºC)	7	Field, Lake Texoma (Oklahoma, Work and Gophen 1999). Menidia audens probably tolerates lower temperatures.
Maximum Temperature (ºC)	35	CTM (Critical Thermal Maximum),gradual temperatue increase, at 16 PSU, 33.5 in at 0 PSU (Davis et al. 2019)
Minimum Salinity (‰)	0	This is a freshwater species.
Maximum Salinity (‰)	50	Menidia audens was found in South San Francisco Bay salt ponds with mean salinities of ~25-50 PSU (Mejia et al. 2008). Fish from Lake Texoma, Oklahoma, had 70% survival for 24 hours at 33 PSU NaCl with direct transfer. Survival of controls was poor, so the experimental period was shore (Hubbs et al. 1971).
Minimum Reproductive Temperature	13.2	Experimental, Hubbs et al. 1971
Maximum Reproductive Temperature	34.2	None
Minimum Reproductive Salinity	0	This is a freshwater species.
Maximum Reproductive Salinity	9	Experimental, Hubbs et al. 1971
Minimum Length (mm)	50	Mense 1967
Maximum Length (mm)	130	Mense 1957, Boomer Lake, OK, more usually 100-119 (Mense 1957; Suttkus 2005)
Broad Temperature Range	None	Warm-Temperate
Broad Salinity Range	None	Nontidal Limnetic-Hyperhaline

General Impacts

Mississippi Silverside (Menidia audens) was introduced to Clear Lake, California for biocontrol of midge larvae without much evaluation of their feeding and effectiveness. They rapidly became vary abundant and spread downriver to the San Francisco estuary. In Clear Lake, they contributed to the extinction of the Clear Lake Splittail (Pogonichthys ciscoides), and in the estuary they are a competitor and predator to the endangered Delta Smelt (Hypomessus transpacificus) (Moyle 2002; Bennet 2005; Baerwald et all. 2012; Schreier et al. 2016). When Mississippi Silversides colonized reservoirs in Oklahoma. Tennessee, and Alabama they led to declines of the native Brook Silverside (Labidesthes sicculus) (Mense 1967; Strongin et al. 2011; Simmons 2013).

Economic Impacts

In California Mississippi Silversides (Menidia audens) were introduced to the Upper and Lower Blue Lakes, and Clear Lake for the control of the Clear Lake Gnat (Chaoborus astictopus). The release in Clear Lake was not authorized by the California Fish and Game Commission, and the effectiveness of the fish for biocontrol was not evaluated (Dill and Cordone 1997; Moyle 2002). The fish was not successful for biocontrol, but became an abundant forage fish for gamefishes. Mississippi Silversides do feed on midge larvae when abundant, but feed mostly in the littoral zone on a wide variety of prey, while midges are abundant throughout the lake (Dill and Cordone 1997; Moyle 2002). The benefits of the fish as a forage fish in Clear Lake are unclear, since they may have largely replaced native minnows without increasing prey biomass (Moyle 2002; Moyle and Holzhauser 1978).

Ecological Impacts

Mississippi Silversides (Menidia audens) have had major ecological impacts in the San Francisco Bay watershed and estuary. In Clear Lake, they are believed to have contributed to the extinction of the endemic Clear Lake Splittail (Pogonichthys ciscoides), probably through competition (Moyle 2002). In the Delta, they are competitors and predators on the endangered Delta Smelt (Hypomesus transpacifius) (Bennet 2005; Baerwald et all. 2012; Schreier et al. 2016). They are an important prey item for major predatory fishes in the estuary, but their increase over the last four decades has not offset an overall decrease in plankton, planktivorous fishes, and larger predatory fishes, summarized under the rubric 'Pelagic Organism Decline', and attributed to large-scale changes in the Bay's hydrology and foodweb (Mac Nally et al. 2010).

In reservoirs in the Mississippi basin (Oklahoma, Alabama, and Tennessee) the range expansion or invasion of Mississippi Silversides has led to explosive colonization. This in turn has resulted in a decline, and sometimes disappearance, of the native Brook Silverside (Labidesthes sicculus) (Mense 1967; Strongin et al. 2011; Simmons 2013). The Mississippi Silverside has a broad diet that significantly overlaps with the Brook Silverside. It also feeds heavily on an abundant, non-native cladoceran, Daphnia lumholtzi, which the Brook Silverisde avoids (Strongin et al. 2011).

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Regional Impacts

NEP-V	Northern California to Mid Channel Islands		Ecological Impact	Predation
DNA surveys of Mississippi Silversides have detected DNA from the endangered, endemic Delta Smelt (Hypomesus transpacificus). In one survey, 41% of the 37 Mississippi Silversides caught by trawling had DNA of Delta Smelt in their guts, but none of the 614 silversides caught by beach seining had traces of Delta Smelt, suggesting that the Smelt were using the deeper, channel habitat (Baerwald et al. 2012). In a later study sampling multiple habitats by multiple methods, 69 of 550 Mississippi Silversides tested positive for Delta Smelt DNA. Predation was more common at sites with clearer water. Reduced water turbidity has favored increased predation on Delta Smelt (Schreier et al. 2018).
P090	San Francisco Bay		Ecological Impact	Predation
DNA surveys of Mississippi Silversides have detected DNA from the endangered, endemic Delta Smelt (Hypomesus transpacificus). In one survey, 41% of the 37 Mississippi Silversides caught by trawling had DNA of Delta Smelt in their guts, but none of the 614 silversides caught by beach seining had traces of Delta Smelt, suggesting that the Smelt were using the deeper, channel habitat (Baerwald et al. 2012). In a later study, sampling multiple habitats by multiple methods, 69 of 550 Mississippi Silversides tested positive for Delta Smelt DNA. Predation was more common at sites with clearer water. Reduced water turbidity has favored increased predation on Delta Smelt (Schreier et al. 2018).
P090	San Francisco Bay		Ecological Impact	Competition
Mississippi Silversides have a broad diet, including zooplankton, benthic invertebrates, and some fish. While Delta Smelt are primarily planktivores, Nonetheless, the two species have a substantial niche overlap. In mixed species aquarium experiments, both species fed on brine shrimp (Artemia sp.) nauplii, but 20 Silversides grew more than twice as fast as Delta Smelt, with 100% survival, while 6 Delta Smelt died These experiments must be applied with caution to the field but suggest the potential for competition between the species (Bennett 2005).
P090	San Francisco Bay		Ecological Impact	Food/Prey
Mississippi Silversides are a frequent prey of such predatory species of Striped Bass (Morone saxatilis), Largemouth Bass (Micropterus salmoides), and the native Sacramento Pikeminnow (Ptychocheilus grandis) (Nobriga and Feyrer 2008). However, Mississippi Silversides have not increased the abundance of predatory fishes, and are part of the overall phenomenon of 'Pelagic Organism Decline' in San Francisco Bay (Mac Nally et al. 2010).
NEP-V	Northern California to Mid Channel Islands		Ecological Impact	Competition
Mississippi Silversides have a broad diet, including zooplankton, benthic invertebrates, and some fish. While Delta Smelt are primarily planktivores. Nonetheless, the two species have a substantial niche overlap. In mixed species aquarium experiments, both species fed on brine shrimp (Artemia sp.) nauplii, but 20 Silversides grew more twice as fast as Delta Smelt, with 100% survival, while six Delta Smelt died. These lab experiments must be applied with caution but suggest the potential for competition between the species (Bennett 2005).
NEP-V	Northern California to Mid Channel Islands		Ecological Impact	Food/Prey
Mississippi Silversides are a frequent prey of such predatory species of Striped Bass (Morone saxatilis), Largemouth Bass (Micropterus salmoides), and the native Sacramento Pikeminnow (Ptychocheilus grandis) (Nobriga and Feyrer 2008). However, Mississippi Silversides have not increased the abundance of predatory fishes, and are part of the overall phenomenon of 'Pelagic Organism Decline' in San Francisco Bay.
CA	California		Ecological Impact	Competition
Mississippi Silversides have a broad diet, including zooplankton, benthic invertebrates, and some fish. While Delta Smelt are primarily planktivores. Nonetheless, the two species have a substantial niche overlap. In mixed species aquarium experiments, both species fed on brine shrimp (Artemia sp.) nauplii, but 20 Silversides grew more twice as fast as Delta Smelt, with 100% survival, while six Delta Smelt died. These lab experiments must be applied with caution but suggest the potential for competition between the species (Bennett 2005)., Mississippi Silversides have a broad diet, including zooplankton, benthic invertebrates, and some fish. While Delta Smelt are primarily planktivores, Nonetheless, the two species have a substantial niche overlap. In mixed species aquarium experiments, both species fed on brine shrimp (Artemia sp.) nauplii, but 20 Silversides grew more than twice as fast as Delta Smelt, with 100% survival, while 6 Delta Smelt died These experiments must be applied with caution to the field but suggest the potential for competition between the species (Bennett 2005).
CA	California		Ecological Impact	Food/Prey
Mississippi Silversides are a frequent prey of such predatory species of Striped Bass (Morone saxatilis), Largemouth Bass (Micropterus salmoides), and the native Sacramento Pikeminnow (Ptychocheilus grandis) (Nobriga and Feyrer 2008). However, Mississippi Silversides have not increased the abundance of predatory fishes, and are part of the overall phenomenon of 'Pelagic Organism Decline' in San Francisco Bay., Mississippi Silversides are a frequent prey of such predatory species of Striped Bass (Morone saxatilis), Largemouth Bass (Micropterus salmoides), and the native Sacramento Pikeminnow (Ptychocheilus grandis) (Nobriga and Feyrer 2008). However, Mississippi Silversides have not increased the abundance of predatory fishes, and are part of the overall phenomenon of 'Pelagic Organism Decline' in San Francisco Bay (Mac Nally et al. 2010).
CA	California		Ecological Impact	Predation
DNA surveys of Mississippi Silversides have detected DNA from the endangered, endemic Delta Smelt (Hypomesus transpacificus). In one survey, 41% of the 37 Mississippi Silversides caught by trawling had DNA of Delta Smelt in their guts, but none of the 614 silversides caught by beach seining had traces of Delta Smelt, suggesting that the Smelt were using the deeper, channel habitat (Baerwald et al. 2012). In a later study sampling multiple habitats by multiple methods, 69 of 550 Mississippi Silversides tested positive for Delta Smelt DNA. Predation was more common at sites with clearer water. Reduced water turbidity has favored increased predation on Delta Smelt (Schreier et al. 2018)., DNA surveys of Mississippi Silversides have detected DNA from the endangered, endemic Delta Smelt (Hypomesus transpacificus). In one survey, 41% of the 37 Mississippi Silversides caught by trawling had DNA of Delta Smelt in their guts, but none of the 614 silversides caught by beach seining had traces of Delta Smelt, suggesting that the Smelt were using the deeper, channel habitat (Baerwald et al. 2012). In a later study, sampling multiple habitats by multiple methods, 69 of 550 Mississippi Silversides tested positive for Delta Smelt DNA. Predation was more common at sites with clearer water. Reduced water turbidity has favored increased predation on Delta Smelt (Schreier et al. 2018).

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