Fishes of Wisconsin
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Round Goby
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SYSTEMATICS and
TAXONOMY
DESCRIPTION
Morphometry and
meristics
Pigmentation
Similar species
Size, growth and age
DISTRIBUTION,
STATUS and HABITAT
Matthew S. Kornis and Norman Mercado-Silva, Univerisity of Wisconsin-Madison
BIOLOGY
Publication Date - April 2011
Reproduction and
Spawning
Ecology
Diet
Associated Species
Importance and
Management
Common
Name:
Round Goby
Other
Common
Names:
Azov goby, Caspian goby, black spotted
goby, and many others in their native
range (Charlebois et al. 1997)
Scientific
name:
Neogobius melanostomus
Wisconsin Apollonia melanostoma (Stepien and
Synonyms: Tumeo 2006)
Etymology: Neogobius - Greek neos, meaning new;
Latin gobius, meaning gudgeon or small
fish
melanostomus - Greek melanos,
meaning black; Greek stoma, meaning
mouth
SYSTEMATICS AND TAXONOMY:
The round goby is a member of the Gobiidae, one of most diverse fish families in the
world but one not native to Wisconsin. The round goby was first described by Pallas
(1811) from specimens taken from the Black Sea at Balaklava, Sevastopol, in the
southern Ukraine on the Crimean peninsula. Two subspecies have been recognized,
one from the Caspian Sea (Neogobius melanostomus affinis) and the other from the
Black Sea (Neogobius melanostomus melanostomus) (Berg 1965). The subspecies
are distinguishable based on the number of transverse scale rows (48-49 in N. m.
melanostomus; 45-46 in N. m. affinis) (Berg 1965). The form found in North America
is N. m. melanostomus. Some systematic and taxonomic studies (Tsyplakov 1974,
cited in Charlebois et al. 1997; Stepien and Tumeo 2006) have proposed that round
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goby be assigned to another genus, Apollonia, and that Neogobius be switched to a
subgenus within Apollonia (i.e., Apollonia (Neogobius) melanostoma).
hybrids involving round goby and the monkey goby (N. fluviatilis) have been reported
from the native range of the round goby, but the monkey goby is limited to Eurasia,
and no round goby hybrids have been reported from North America (Charlebois et al.
1997).
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DESCRIPTION:
Morphometry and meristics:
The quantitative data presented here are from the round goby's native range based
on Berg (1965) and Miller (1986) and from two specimens from Lake Superior
(UWMZ 11187) and 48 from Lake Michigan (UWMZ 11046, 11664) (N. Delventhal
and J. Lyons, University of Wisconsin Zoological Museum, personal communication,
2008).
The round goby has an elongated body that is round in cross section. The body
depth goes into standard length (SL) on average 4.3 times (range 3.53-5.15). Head
length goes into SL 3.0-3.6 times and into total length (TL) 4.2-4.5 times. Head depth
goes into head width 0.9-1.2 times. The interorbital width is 0.8-0.9 times the eye
diameter, whereas snout length is 1.1-1.4 times the eye diameter. The mouth is
terminal to slightly subterminal with thick lips, the upper lip narrowing slightly towards
the rear.
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The posterior angle of the jaws extends to below the anterior quarter of the eye. The
tongue is slightly notched. The caudal peduncle depth is about two thirds of the
caudal peduncle length. First dorsal fin height goes into SL 5.6-7.5 times, pectoral fin
length goes into SL 3.5-4.3 times, and pelvic fin disk length goes into SL 4.2-5.3
times.
Sexual dimorphism in morphology is marked in round goby; males have a larger size
at age, enlarged cheeks, and darker coloration (Miller 1984). Both sexes have an
erectile urogenital papilla between the vent and the base of the anal fin. The female
papilla is broad and blunt (0.3-0.5 mm wide, 0.2-0.4 mm long), whereas the male
papilla is longer (0.3-0.6 mm), pointed, and has a terminal notch (Charlebois et al.
1997).
The round goby has two clearly separated dorsal fins. In its native range, the first
dorsal fin usually has seven to eight spines, and the second has one spine and 1217 rays. However, among 50 Lake Superior and Lake Michigan specimens, the first
dorsal fin almost always had only six spines (49 specimens with six spines and one
specimen with five spines), and the second had one spine and 15-17 rays. The anal
fin has one spine and 9-14 rays in the native range and one spine and 11-15 rays in
50 Lake Michigan and Lake Superior specimens. In both the native range and the
Great Lakes, the two pelvic fins are united to form a disc that has 17-20 total rays
(always 20 in 50 Lake Michigan and Lake Superior specimens) and extends to or
almost to the vent.
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This disk can be used to generate suction and allows the round goby to adhere to
solid surfaces. There are 31-34 vertebrae. A lateral line is absent on the flanks, but
some cephalic sensory pores may occur on the head. There are 42-59 scales in the
lateral series. The back, neck, belly, flank, and a portion of the gill covers are
covered by ctenoid scales (Charlebois et al. 1997). In its native range, the top of the
head is reportedly covered by cycloid scales (Stranai and Andreji 2004), but at least
some Great Lakes individuals have ctenoid scales on the top of the head (Kornis,
personal observation). Two pairs of plates of pharyngeal teeth adjoin the dorsal and
ventral portions of the first two branchial arches (four total plates) and are covered
with incisors (Pinchuk 1991; Ghedotti et al. 1995; L. Garrison, Northern Michigan
University, personal communication, 2008).
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Pigmentation:
The round goby has a brownish or yellow-green body with large dark brown lateral
spots. The head is usually darker than the rest of the body. The fins are generally
dark grey, but a large oblong black spot is usually present on the posterior portion of
the first dorsal fin, starting on the fifth ray from the front.
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In breeding males, the body and fins may be almost completely black, with the
pectoral fins sometimes fringed with white or yellow.
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Similar species:
The round goby is most similar to another nonnative species, the tubenose goby.
The tubenose goby has elongated anterior nostrils extending over the upper lip that
the round goby lacks. Conversely, the round goby has a dark blotch on the posterior
portion of the first dorsal fin that is absent in the tubenose goby. The tubenose goby
has four gill rakers on the first gill arch and 24 total pelvic fin rays versus 12 gill
rakers and 20 total pelvic fin rays in the round goby (Miller 1986; Jude et al. 1992).
The round goby is also similar to the native mottled, slimy, spoonhead, and
deepwater sculpins. Unlike the sculpins, which have two distinctly separate pelvic
fins, round goby have pelvic fins that are fused underneath the body to form a disc.
Round goby also have scales on the body, whereas the sculpins lack scales. The
sculpins have one or more spines on the preopercle, and round goby have none.
Further information on identifying the round goby can be found at the website
http://wiscfish.org/.
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Size, growth and age:
The round goby is a relatively small and short-lived species. In its native range,
where it is primarily a marine or brackish-water species, the average adult SL is
about 127 mm, and the maximum SL is about 250 mm (Berg 1965; Charlebois
2002). The maximum lifespan is usually 4-5 years (MacInnis and Corkum 2000a).
Males generally grow faster and achieve larger sizes than females. Males typically
reach 115 mm SL after their first year (range 100-130 mm) and 170-180 mm by their
third or fourth year (Berg 1965; Jude et al. 1992). Conversely, females average 95
mm SL (80-110 mm) after the first year and 130-140 mm by their third or fourth year.
Annual growth increments decrease with age for females and increase with age for
males (Skazkina and Kostyuchenko 1968, cited in Charlebois et al. 1997). Females
normally mature at age 2 to age 3 and males at age 3 to age 4.
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In the Great Lakes, round goby grow more slowly, reach a smaller maximum size,
have a shorter lifespan, and mature earlier than in their native range, though males
still grow faster than females (MacInnis and Corkum 2000a; Corkum et al. 2004). In
the Great Lakes, the average adult SL is about 60-80 mm, and the maximum TL is
180 mm (Jude et al. 1992; MacInnis and Corkum 2000a). Maximum sizes recorded
for Wisconsin waters are 165 mm TL (Kornis, unpublished data) and 131 mm SL (J.
Lyons, Wisconsin DNR, personal communication, 2008). The maximum reported
lifespan in the Great Lakes is five years (French and Black 2009), but few fish
exceed three years in age. Females mature at age 1 to age 2 and males at age 2 to
age 3 (Jude et al. 1992; Charlebois 2002). Growth rates appear to vary within the
Great Lakes, although we know of only two published studies on size at age (Table
1).
Table 1. Mean back-calculated standard length at age from otoliths for round goby
from the Detroit River, Ontario (lapilli otoliths, MacInnis and Corkum 2000a), and
Lake Huron, Michigan (sagittal otoliths, French and Black 2009). Sample sizes (N)
represent the number of round goby captured at a given age in the Detroit River and
the number of back-calculated estimates for Lake Huron (e.g., an age-3 fish is
counted in the sample size (N) for ages 1, 2, and 3 for Lake Huron but is only
counted as age 3 for the Detroit River).
Location
Detroit
River
Detroit
River
Lake
Huron
Size at age (mm)
Sex
1
2
3
4
5
Female
58.4 (N =
99)
62.8 (N =
92)
35.3 (N =
30)
64.6 (N =
44)
75.9 (N =
30)
55.3 (N =
30)
82.7 (N =
3)
-
-
-
-
-
68.0 (N =
26)
77.4 (N =
26)
85.9 (N =
8)
Male
Male
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DISTRIBUTION, STATUS AND HABITAT:
The round goby is native to the Ponto-Caspian region of Eurasia. It is known from
the Azov, Caspian, and Black seas and the lower reaches of large tributary rivers
(e.g., Danube, Dniester, Volga) in Bulgaria, Romania, Russia, Kazakhstan, and
Azerbaijan (Maitland 1977; Miller 1986, cited in Charlebois et al. 1997).
Round goby have recently become established in other parts of Europe and in North
America. They have presumably spread via transport in the ballast water of ships;
the discovery of larval round goby in the water column supports this idea (Hensler
and Jude 2007). In Europe, they have invaded the Gulf of Gdansk (Baltic Sea) and
the upper Danube River (Skora and Stolarski 1993; Stepien and Tumeo 2006). In
North America, round goby were first observed in the St. Clair River and Lake Huron
in Michigan in 1990-91 (Jude et al. 1992; Schaeffer et al. 2005), and by 1995 they
were found in all of the Great Lakes (Charlebois et al. 2001; Clapp et al. 2001;
Corkum et al. 2004). The sources of the first colonists were probably the northern
Black Sea (Dnieper River mouth near Kherson, Ukraine) and the Gulf of Gdansk
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(Charlebois et al. 1997; Dillon and Stepien 2001; J .E. Brown, University of Toledo,
personal communication, 2008), although high levels of genetic diversity in the Great
Lakes suggests that round goby invaded from multiple source populations (Stepien
et al. 2005). Their rapid spread among and within the Great Lakes was again
presumably facilitated via transport in the ballast water of ships, though the presence
of larval round goby near surface waters suggests that currents may also contribute
to their spread within the Great Lakes (Hensler and Jude 2007). Round goby have
entered the Illinois Waterway system in Chicago that connects Lake Michigan to the
Mississippi River basin, and they have migrated downstream quickly, with a few
individuals found more than 190 km inland from Chicago (Irons et al. 2006).
In Wisconsin, round goby are known from specific coastal areas of Lake Michigan
and Lake Superior and the lower reaches of some tributaries.
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They appear to be increasing in abundance and spreading. In Lake Superior, round
goby are found in Superior Harbor and the St. Louis River up to the Fond du Lac
Dam, located over 30 km from the open lake. They have also been observed in the
lower 6.4 km of Amnicon River, Douglas County, which enters Lake Superior 13 km
east of Superior Harbor (D. Pratt, Wisconsin DNR, personal communication, 2008).
In Lake Michigan, round goby are widespread and common in the vicinity of Door
County, particularly in Green Bay and Sturgeon Bay and to a lesser extent in Lake
Michigan proper. Further south, round goby also occur in and around the ports of
Kewaunee, Manitowoc, Sheboygan, Port Washington, Milwaukee, Racine, and
Kenosha (P. Peeters, S. Hogler, T. Burzynski, Wisconsin DNR, personal
communication, 2008). Round goby have been collected in the lower reaches of 26
Green Bay and Lake Michigan tributaries up to 33.9 km upstream of the open lake
(Table 2). In some of these tributaries round goby upstream movements have been
blocked by impassable dams. Habitat models indicate that there are long stretches of
riverine habitat suitable for round goby upstream of these dams, suggesting that if
the dams were no longer barriers the distribution of round goby would expand
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substantially (Kornis and Vander Zanden 2010). Further details on the distribution of
round in Wisconsin can be found at the website
http://infotrek.er.usgs.gov/wdnrfish/map/index.
Table 2. Streams and rivers invaded by round goby in the Lake Michigan basin of
Wisconsin with stream distances from Lake Michigan (from Kornis and Vander
Zanden 2010). Round goby were found immediately downstream of a physical
barrier (dam or waterfall) in the Oconto, East Twin, Suamico, Ahnapee, Kewaunee,
West Twin, and Menominee rivers.
Stream Name
Distance Upstream (km)
Pensaukee River
33.92
Oconto River
23.19
Fox River
19.5
Little Suamico River
17.55
East Twin River
16.59
Suamico River
13.28
Ahnapee River
12.6
Kewaunee River
11.4
Sheboygan River
10.5
West Twin River
9.6
Duck Creek
6.4
Milwaukee River
5.5
Big Creek
5
Manitowoc River
4.7
Little River
4.2
Pigeon River
3.2
Menomonee River
3
Menominee River
2.9
Pike River
2.67
Whitefish Bay Creek
1.6
Root River
0.5
Mud Lake Outlet
0.1
Little Manitowoc River
0.1
Silver Creek
0.1
Stony Creek
0.1
Peshtigo River
0.1
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Round goby can live in a variety of habitats, including freshwater rivers and lakes
and brackish-water areas of marine seas.
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They are bottom dwellers and prefer rocky areas (Jude et al. 1995; Charlebois et al.
1997; Corkum et al. 2004), although they can also be found on sandy substrates
and in vegetated areas (Charlebois 2002) especially during daylight (Ray and
Corkum 2001). In rivers they prefer rocky riffles (Carman et al. 2006) but can be
found in slower moving water as well (Kornis, personal observation). Mainly juveniles
are found in sandy areas, perhaps as a result of exclusion by adults from preferred
rocky habitats. Round goby also use artificial substrates such as pipes and
shipwrecks (Wickett and Corkum 1998) as habitat. However, thus far they appear to
have avoided coastal wetland habitat, possibly due to low availability of rocky
substrate (Cooper et al. 2007; Young et al. 2010). During spring and summer, round
goby in the Great Lakes can be found near shore in depths as shallow as less than 1
m, but in fall and winter they move to deeper waters (Charlebois 2002) and occupy
depths up to 50-60 m (Miller 1986). They may also leave some stream habitats to
over-winter in the Great Lakes (Pennuto et al. 2010).
The abundance of round goby in the Great Lakes has increased rapidly, and they
are now one of the most abundant benthic species (Jude 1997, 2001). In appropriate
habitat, round goby typically occur at densities of 2-6 fish/m 2 with a maximum of 19
fish/m 2 (Ray and Corkum 2001). Individual round goby have a home range of at
least 5 (±1.2) m 2 (Ray and Corkum 2001), but quickly growing goby populations
create high densities that cause competition for space and food and promote range
expansion (Steingraeber and Thiel 2000).
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BIOLOGY:
Reproduction and Spawning:
Round goby have a relatively long spawning season. Egg development begins in
September in preparation for spawning the following spring (Kulikova 1985). In
spring, round goby begin moving into shallow waters to spawn when water
temperatures exceed 9°C, with males preceding females (Charlebois et al. 1997).
Spawning depths are generally 0.2-1.5 m (MacInnis and Corkum 2000b) but may be
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as deep as 7-11 m (Wickett and Corkum 1998). In their native range, round goby
spawning occurs between May and July (Maitland 1977) when temperatures exceed
12°C (Moiseyeva and Rudenko 1996) and can extend into September (Berg 1965).
Similarly, in North America round goby begin to spawn in the spring and continue to
spawn throughout the summer. Spawner densities decline after July, but spawners
can still be found in October (MacInnis and Corkum 2000b).
Males establish a nest in the cavity underneath a hard object. Many objects can
serve as nests with the general requirements being the existence of an immobile
overhead surface and a single opening to the cavity (Miller 1984). Males compete for
nesting territories; larger males have an advantage in holding preferred nest sites,
potentially limiting the number of nests in a given area (Stammler and Corkum 2005).
Males attract females to the nest by producing a call, with receptive females
responding with a quieter call (Charlebois et al. 1997; Rollo et al. 2007; Meunier et
al. 2009). Males also produce a sex pheromone to attract mates, and behavioral
studies indicate that these pheromones may induce spawning behavior in females
(Zielinski et al. 2003; Gammon et al. 2005; Corkum et al. 2006). A response by the
female will induce the male to secrete a sticky secretion with which it coats the
spawning surface of the nest. Males will chose certain females for spawning and
reject others (Meunier et al. 2009). Chosen females enter the nest and lay their eggs
on the overhead surface of the cavity. As each egg is extruded, the extended base of
each egg is glued to the surface (Moskal'kova 1996). Fertilization rates are
approximately 95%. Multiple females may spawn with one male, and as many as
10,000 eggs from four to six females can be deposited in a single nest. A female can
spawn every 20 days and up to six times per year, but usually females produce only
two to four clutches of eggs per season in the wild (MacInnis and Corkum 2000b).
Clutch size increases with female size; a 40 mm SL female can produce
approximately 70 eggs per clutch whereas females of 70 mm and 96 mm can
produce 250 and 600 eggs per clutch, respectively (MacInnis and Corkum 2000b).
Fecundity is positively correlated with standard length in the Detroit River, Ontario
(Number of eggs = -331.89 + (8.95*SL [mm])), where absolute fecundity, the eggs
produced per female per season, can be 252 to 1,818 eggs (MacInnis and Corkum
2000b). These values are in the lower portion of the range (328-5,221 eggs) reported
by Kovtun (1979) for the absolute fecundity of round goby in their native range.
However, round goby in the Great Lakes have higher relative fecundity (i.e., eggs
per gram of fish) than native species (MacInnis and Corkum 2000b).
After spawning, the male round goby guards, maintains, and aerates the eggs in the
nest (Meunier et al. 2009). This contributes to an estimated 95% hatching success
rate (Wickett and Corkum 1998; Charlebois 2002), although larger nests can suffer
higher egg mortality from predation and insufficient aeration (Kovtun 1979). Once the
eggs hatch, the fry remain in or near the nest under the at least partial protection of
the male for four to nine days, after which they disperse. During nest guarding
periods, males do not feed (Charlebois et al. 1997), and most die soon after the
spawning period is finished (Jude et al. 1992). Thus most male round goby only
spawn once in their lifetimes.
Some male round goby have been reported to spawn without establishing or
defending a nest. Instead, they use a "sneaker" or "satellite" spawning strategy
(Gross 1991). The sneaker male mimics a female in size and appearance in order to
deceive the nesting male and gain access to the nest while the nesting male is
spawning with a female. The sneaker male then attempts to fertilize some of the
eggs being spawned. After spawning, the sneaker male quickly leaves the nest
before he can be recognized as a fraud and attacked by the nesting male (MacInnis
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and Corkum 2000b).
A study from the native range of the round goby indicated that the ratio of adult male
to female round goby was inversely related to fry survival (Kovtun 1979). In years
when the ratio was high (up to 1.9 males per female), fry survival was low (6-14%),
but in years when the ratio was low (1.1-1.4 males per female), fry survival was much
higher (70-93%). Low survival when males were relatively numerous may have
occurred because nesting males could not adequately defend eggs in the presence
of many non-nesting males (Charlebois et al. 1997).
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Development:
Round goby eggs are oval with a sharp apex and an approximate mean diameter of
3.2. Eggs have a fiber-like basal stalk 0.4-0.5mm long (Leslie and Timmins 2004). A
relatively high degree of development occurs within the egg. The eyes develop early
and are light sensitive before hatching. Complete gonad development and early
stages of gametogenesis also occur within the egg. Consequently, some authors
have suggested that round goby do not have a true larval stage and hatch as
juveniles (Moskal'kova 1996; but see Miller 1984 and Hensler and Jude 2007). A
complete developmental history for round goby is found in Charlebois et al. (1997) A
description of age-0 round goby is found in Leslie and Timmins (2004)
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Ecology:
Round goby can tolerate a wide range of environmental conditions, an attribute that
has facilitated their invasion and expansion in the Great Lakes via ship ballast water,
which often presents harsh conditions. Round goby can withstand low levels of
dissolved oxygen (0.3-0.9 mg/L), although if given the opportunity, they will leave
areas in which dissolved oxygen is less than 50-60% saturation (Svetovidov 1964,
cited in Charlebois et al. 1997). They occur at a wide variety of salinities, from
freshwater to 40.6 parts per thousand (Kazancheev 1981, cited in Charlebois et al.
1997; full seawater is about 30 parts per thousand), and they can reproduce at
salinities of up to 19.5 parts per thousand (Moskal'kova 1996, cited in Charlebois et
al. 1997). Round goby prefer rocky substrate (Ray and Corkum 2001) but are
adaptable to a variety of other habitats (Charlebois et al. 1997; Ahnelt et al. 1998;
Young et al. 2010). They have a wide range of feeding behaviors including the ability
to feed in total darkness (Jude et al. 1995; Carman et al. 2006). Round goby are
eurythermal, tolerating temperatures between -1°C and 30°C (Moskal'kova 1996).
Their optimum temperature for growth occurs from 23°C to 26°C (Lee and Johnson
2005). Overall, round goby are considered to have relatively high tolerance to
chronic exposure to toxic substances (Spromberg and Birge 2005).
Round goby are preyed upon by many species of fish and other vertebrates (i.e.,
water snakes, birds, etc.). They are eaten by walleye, smallmouth bass, rock bass,
tubenose goby, stonecat, yellow perch, northern pike, freshwater drum, and trout
and salmon (particularly brown trout) (Jude et al. 1995; Campbell et al. 2009; Kornis,
unpublished data from angler interviews). Round goby are found in the diets of
double-crested cormorants to varying degrees depending on location and season
(Somers et al. 2003; Ross et al. 2004). As round goby have become more common,
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a threatened species of water snake in Lake Erie has shifted its diet to primarily
consume round goby (King et al. 2006).
Predation influences the behavior and distribution of round goby. Round goby will
burrow into the substrate to avoid predators (Jude et al. 1992). Smaller individual
round goby are more susceptible to predation than larger ones, and predation rates
are lower in sheltered habitats (Belanger and Corkum 2003). Vulnerability to
predation thus plays a crucial role in round goby habitat selection. High predator
abundance has been hypothesized as a barrier to round goby expansion in the
Shiawassee and Clinton rivers in Michigan and may also be a limiting factor in the
coastal wetlands of Green Bay (D. Jude, University of Michigan, personal
communication, 2008).
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Diet:
Round goby are carnivores with great dietary flexibility. They generally feed on
invertebrates, especially bivalve mollusks (primarily non-native zebra and quagga
(dreissenid) mussels in the Great Lakes), small crustaceans (amphipods), and
aquatic insects (particularly in streams), although larger individuals are known to
consume fish and fish eggs, including their own species (Berg 1965; Charlebois
2002; Steinhart et al. 2004a; Carman et al. 2006; Campbell et al. 2009; Pennuto et
al. 2010; Kornis, unpublished data). Round goby diets may also include snails, softshelled crayfish, and zooplankton (Ray and Corkum 1997; Carman et al. 2006).
Round goby undergo a diet shift beginning at about 50-60 mm SL; at smaller sizes
they typically consume arthropods, but as they grow larger they switch to diets
dominated by mollusks (French and Jude 2001; Janssen and Jude 2001; Jude et al.
2005; Campbell et al. 2009). In a laboratory study, Diggins et al. (2002) found that
round goby have a preference for amphipods over dreissenid mussels in bare
habitats and in habitats with clear visibility, but they will consume more dreissenid
mussels in more complex or lower visibility habitats where amphipods are not as
readily found. In streams round goby will consume both benthic and drifting
invertebrates (Carman et al. 2006; Pennuto et al. 2010).
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Associated Species:
Over their Great Lakes range, round goby co-occur with many different fishes. In
Wisconsin waters, round goby are found in the same microhabitats as the native
benthic species logperch, johnny darter, and slimy and mottled sculpin. At the
macrohabitat scale, they are encountered most commonly with alewife, a variety of
salmonids, northern pike, smallmouth bass, rock bass, a variety of sunfishes, yellow
perch, and walleye (Jude et al. 1995; Steinhart et al. 2004b; Dopazo et al. 2008;
Kornis, personal observation).
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Importance and Management:
The fisheries importance of round goby differs between Eurasia and North America.
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Fishes of Wisconsin
Round goby have considerable value as a food fish in their native range, where they
are marketed fresh or canned (Maitland 1977; Miller 1986; Charlebois et al 1997).
However, their use as food in the Great Lakes is limited by their smaller size and by
regulations restricting possession. In Wisconsin, it is illegal for fishers to possess
round goby except that one may be killed and transported to a WDNR office to
confirm identification. The goal of this regulation is to prevent the use of round goby
as bait or food by anglers and their accidental or intentional spread into inland
waters. Where common, round goby are easy to catch by angling, and they provide
some minor sport, but most anglers consider them a bait-stealing nuisance.
The primary importance of round goby in the Great Lakes is in their influence on
other species and on ecosystem structure and function. The establishment of round
goby throughout the Great Lakes has caused declines in a variety of native species,
altered the structure and function of food webs, and possibly influenced ecosystem
productivity (Kuhns and Berg 1999; Vanderploeg et al. 2002; Lee and Johnson
2005; Campbell et al. 2009). Abundant round goby may reduce benthic invertebrate
populations, which are important food for many native fishes. Round goby density
was found to correlate negatively with the abundance of benthic invertebrates,
including non-native dreissenid mussels and native isopods, amphipods, and snails,
in rocky habitats of the Door Peninsula in Green Bay (Lederer et al. 2006, 2008). The
increase in round goby and dreissenid mussels together was associated with
declines in native invertebrates in southern Lake Michigan (Kuhns and Berg 1999).
Round goby have displaced and reduced populations of native benthic fishes
throughout the Great Lakes, particularly mottled and slimy sculpin, logperch, and
johnny darter, by competing for food and spawning areas and by feeding on their
eggs (Jude et al. 1992, 1995; French and Jude 2001; Janssen and Jude 2001;
Lauer et al. 2004; Balshine et al. 2005). Egg predation by round goby may also
impede the recovery of depressed populations of native lake sturgeon (Nichols et al.
2003; Lee and Johnson 2005) and lake trout (Chotkowski and Marsden 1999).
Round goby quickly consume smallmouth bass eggs if smallmouth bass males
guarding their nests are removed by anglers (Steinhart et al. 2004a).
In certain instances, abundant round goby have provided some benefits to the Great
Lakes ecosystem. Steinhart et al. (2004b) reported that young-of-the-year
smallmouth bass predation on round goby in Lake Erie led to an increase in
smallmouth bass growth rates, which may have improved smallmouth bass survival.
A rare species of water snake in Lake Erie fed heavily on round goby after the fish
became common, and consequently water snake individual growth rate and
populations size increased (King et al. 2006). Round goby predation can reduce the
abundance and alter the size structure of nonnative dreissenid mussels in some
instances (Djuricich and Janssen 2001; Lederer et al. 2006). However, round goby
are not likely to control dreissenid mussel populations fully because they prefer to
feed on smaller mussels, leaving larger mussels to reproduce (Ray and Corkum
1997), and they are unable to consume mussels hidden under rocks (Djuricich and
Janssen 2001).
Round goby have changed the flow of energy, toxins, and diseases in the Great
Lakes ecosystem, generally to the detriment of native species (Kwon et al. 2006). In
areas of Lake Erie where dreissenid mussels have high levels of contaminants and
where round goby feed heavily on the mussels, consumption of round goby by
smallmouth bass may lead to higher-than-expected levels of mercury in the
smallmouth bass (Hogan et al. 2007). High concentrations of polychlorinated
biphenyls (PCBs) have been recorded in round goby in the PCB-contaminated
Raisin River, a tributary to Lake Erie, and consumption of round goby could lead to
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Fishes of Wisconsin
bioaccumulation of PCBs in predators (Jude 2001; Hanari et al. 2004). High
concentrations of other contaminants (i.e., perfluorinated compounds [PFCs]) in
round goby have also been identified in some areas (Kurunthachalam et al. 2005).
Round goby have been suggested as a vector for the transfer of the disease botulism
Clostridium botulinum type E to fish-eating birds in the Great Lakes (Corkum et al.
2004; Yule et al. 2006). Movement of infected round goby via ballast water could
potentially expand the range of the newly observed fish disease viral hemorrhagic
septicemia (VHS) (Egan 2006). This disease has recently caused mass die-offs of
gizzard shad, muskellunge, lake whitefish, yellow perch, walleye, freshwater drum,
and round goby in the eastern Great Lakes (Whelan 2007). Finally, there has been
speculation that consumption of dreissenid mussels and other invertebrates by
abundant round goby may change the cycling and availability of phosphorus, a
limiting nutrient for algae and plants in the Great Lakes and ultimately change
patterns of benthic primary and secondary production. However, ecological modeling
suggests that despite being quite common in Lake Erie, round goby apparently have
had relatively little effect on phosphorus cycling (Bunnell et al. 2005).
Once established, round goby are nearly impossible to eliminate, so round goby
management has focused on preventing their spread and to a lesser extent limiting
their abundance where they already occur. Of primary concern has been the
movement of round goby from the Great Lakes basin to the Mississippi River basin
via the Illinois Waterway system. An electrical barrier has been installed near
Chicago to block the passage of round goby and other nonnative fish species
through the waterway (Charlebois 2002). The long-term effectiveness of this barrier
is uncertain. A complimentary strategy to prevent the expansion of round goby
beyond the Great Lakes includes coordinated regulations and education efforts
among the Great Lakes states and the Province of Ontario banning the use of round
goby as bait and their possession by anglers. There are also efforts to better
regulate ballast water discharges from Great Lakes shipping. Ships bound for the
Great Lakes are required to flush their ballast tanks with seawater prior to entering
the St. Lawrence Seaway, must submit documentation of compliance with ballast
regulations, and are privy to inspection (99% of Great Lakes bound vessels received
ballast tank inspections in 2008, with 98.6% of tanks in compliance with current
regulations) (Great Lakes Seaway Ballast Water Working Group 2009).
Efforts to limit round goby abundance have thus far focused on enhancing predator
numbers and consumption through more restrictive angling regulations and in some
cases stocking. For example, in Superior Harbor and associated bays and tributaries
in northwestern Wisconsin, angler bag limits for northern pike and walleye were
reduced in response to the establishment of several nonnative invasive fishes
including ruffe, white perch, round goby, and tubenose goby. Some scientists have
proposed controlling round goby numbers by utilizing round goby male sex
pheromones to disrupt reproduction and to concentrate round goby females for
removal (Corkum et al. 2006). Sex pheromones have yet to be employed in the
control of invasive fishes, but they have been used extensively in the control and
eradication of insect pests (Li et al. 2003).
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Citations:
Ahnelt, H., P. Banarescu, R. Spolwind, A. Harka, and H. Waidbacher. 1998.
Occurrence and distribution of three gobiid species (Pisces, Gobiidae) in the
middle and upper Danube region - examples of different dispersal patterns?
http://infotrek.er.usgs.gov/wdnr_fishes/account.jsp?species_param=1490[7/10/2012 2:03:29 PM]
Fishes of Wisconsin
Biologia 53:665-678.
Balshine, S., A. Verma, V. Chant, and T. Theysmeyer. 2005. Competitive
interactions between round gobies and logperch. Journal of Great Lakes
Research 31:68-77.
Belanger, R. M., and L. D. Corkum. 2003. Susceptibility of tethered round gobies
(Neogobius melanostomus) to predation in habitats with and without shelters.
Journal of Great Lakes Research 29:588-593.
Berg, L. S. 1965. Freshwater fishes of the U.S.S.R. and adjacent countries.
Translated by Israel Program for Scientific Translations Ltd., Jerusalem.
Translated from original Russian version (1949), Academy of Sciences U.S.S.R.
Zoological Institute.
Bunnell, D. B., T. B. Johnson, and C. T. Knight. 2005. The impact of introduced
round gobies (Neogobius melanostomus) on phosphorus cycling in central Lake
Erie. Canadian Journal of Fisheries and Aquatic Sciences 62:15-29.
Campbell, L. M., R. Thacker, D. Barton, D. C. G. Muir, D. Greenwood, and R. E.
Hecky. 2009. Re-engineering the eastern Lake Erie littoral food web: the trophic
function of non-indigenous Ponto-Caspian species. Journal of Great Lakes
Research 35:224-231.
Carman, S. M., J. Janssen, D. J. Jude, and M. B. Berg. 2006. Diel interactions
between prey behaviour and feeding in an invasive fish, the round goby, in a
North American river. Freshwater Biology 51:742-755.
Charlebois, P. M. 2002. The round goby (Neogobius melanostomus pallas). Lakeline
22:46-47.
Charlebois, P. M., J. E. Marsden, R. G. Goettel, R. K. Wolfe, D. J. Jude, and S.
Rudnika. 1997. The round goby Neogobius melanostomus (Pallas): a review of
European and North American literature. Illinois-Indiana Sea Grant Program and
Illinois Natural History Survey, INHS Special Publication 20, Champaign.
Charlebois, P. M., L. D. Corkum, D. J. Jude, and C. Knight. 2001. The round goby
(Neogobius melanostomus) invasion: current research and future needs. Journal
of Great Lakes Research 27:263-266.
Chotkowski, M. A., and J. E. Marsden. 1999. Round goby and mottled sculpin
predation on lake trout eggs and fry: field predictions from laboratory experiments.
Journal of Great Lakes Research 25:26-35.
Clapp, D. F., P. J. Schneeberger, D. J. Jude, G. Madison, and C. Pistis. 2001.
Monitoring round goby (Neogobius melanostomus) population expansion in
eastern and northern Lake Michigan. Journal of Great Lakes Research 27:335341.
Cooper, M. J., C. R. Ruetz III, D. G. Uzarski, and T. M. Burton. 2007. Distribution of
round gobies in coastal areas of Lake Michigan: are wetlands resistant to
invasion? Journal of Great Lakes Research 33:303-313.
Corkum, L. D., M. R. Sapota, and K. E. Skora. 2004. The round goby, Neogobius
melanostomus, a fish invader on both sides of the Atlantic Ocean. Biological
Invasions 6:173-181.
Corkum, L. D., W. J. Arbuckle, A. J. Belanger, D. B. Gammon, W. Li, A. P. Scott,
and B. Zielinski. 2006. Evidence of a male sex pheromone in the round goby
(Neogobius melanostomus). Biological Invasions 8:105-112.
Diggins, T. P., J. Kaur, R. K. Chakraborti, and J. V. DePinto. 2002. Diet choice by
the exotic round goby (Neogobius melanostomus) as influenced by prey mortality
and environmental complexity. Journal of Great Lakes Research 28:411-420.
http://infotrek.er.usgs.gov/wdnr_fishes/account.jsp?species_param=1490[7/10/2012 2:03:29 PM]
Fishes of Wisconsin
Dillon, A. K., and C. A. Stepien. 2001. Genetic and biogeographic relationships of
the invasive round (Neogobius melanostomus) and tubenose (Proterorhinus
marmoratus) gobies in the Great Lakes versus Eurasian populations. Journal of
Great Lakes Research 27:267-280.
Djuricich, P., and J. Janssen. 2001. Impact of round goby predation on zebra
mussel size distribution at Calumet Harbor, Lake Michigan. Journal of Great
Lakes Research 27:312-318.
Dopazo, S. N., L. D. Corkum, and N. E. Mandrak. 2008. Fish assemblages and
environmental variables associated with gobiids in nearshore areas of the lower
Great Lakes. Journal of Great Lakes Research 34:450-460.
Egan, D. 2006. Fish virus could limit shipping: Michigan seeks ban on ballast from
eastern Great Lakes. Milwaukee Journal Sentinel (November 19)
French, J. R. P., III, and D. J. Jude. 2001. Diets and diet overlap of nonindigenous
gobies and small benthic native fishes co-inhabiting the St. Clair River, Michigan.
Journal of Great Lakes Research 27:300-311.
French, J. R. P., III, and M. G. Black. 2009. Maximum length and age of round
gobies (Apollonia melanostomus) in Lake Huron. Journal of Freshwater Ecology
24:173-175.
Gammon, D. B., W. Li, A. P. Scott, B. S. Zielinski, and L. D. Corkum. 2005.
Behavioural responses of female Neogobius melanostomus to odours of
conspecifics. Journal of Fish Biology 67:615-626.
Ghedotti, M. J., J. C. Smihula, and G. R. Smith. 1995. Zebra mussel predation by
round gobies in the laboratory. Journal of Great Lakes Research 21:665-669.
Great Lakes Seaway Ballast Water Working Group. 2009. 2008 Summary of Great
Lakes Seaway Ballast Water Working Group. Available:
http://www.d9publicaffairs.com/posted/443/Document.261306.pdf.
Gross, M. R. 1991. Evolution of alternative reproductive strategies: frequency
dependent sexual selection in male bluegill sunfish. Philosophical Transactions of
the Royal Society of London Series B: Biological Sciences 332:59-66.
Hanari, N., K. Kannan, Y. Horii, S. Taniyasu, N. Yamashita, D. J. Jude, and M. B.
Berg. 2004. Polychlorinated naphthalenes and polychlorinated biphenyls in
benthic organisms of the Great Lakes food chain. Archives of Environmental
Contamination and Toxicology 47:84-93.
Hensler, S. R., and Jude, D. J. 2007. Diel vertical migration of round goby larvae in
the Great Lakes. Journal of Great Lakes Research 33:295-302.
Hogan, L. S., E. Marschall, C. Folt, and R. A. Stein. 2007. How non-native species
in Lake Erie influence trophic transfer of mercury and lead to top predators.
Journal of Great Lakes Research 33:46-61.
Irons, K. S., M. A. McClelland, and M. A. Pegg. 2006. Expansion of round goby in
the Illinois Waterway. American Midland Naturalist 156:198-200.
Janssen, J., and D. J. Jude. 2001. Recruitment failure of mottled sculpin Cottus
bairdi in Calumet Harbor, southern Lake Michigan, induced by the newly
introduced round goby Neogobius melanostomus. Journal of Great Lakes
Research 27:319-328.
Jude, D. J. 1997. Round gobies: cyberfish of the third millennium. Great Lakes
Research Review 3:27-34.
Jude, D. J. 2001. Round and tubenose gobies: 10 years with the latest Great Lakes
phantom menace. Dreissena 11:1-14.
Jude, D. J., D. Albert, D. G. Uzarski, and J. Brazner. 2005. Lake Michigan's coastal
http://infotrek.er.usgs.gov/wdnr_fishes/account.jsp?species_param=1490[7/10/2012 2:03:29 PM]
Fishes of Wisconsin
wetlands: distribution, biological components with emphasis on fish and threats.
Pages 439-477 in M. Munawar and T. Edsall, editors. The state of Lake Michigan:
ecology, health and management. Ecovision World Monograph Series, SPB
Academic Publishing, Amsterdam.
Jude, D. J., J. Janssen, and G. Crawford. 1995. Ecology, distribution, and impact of
the newly introduced round and tubenose gobies on the biota of the St. Clair and
Detroit Rivers. Pages 447-460 in M. Munawar, T. Edsall and J. Leach, editors.
The Lake Huron ecosystem: ecology, fisheries, and management. Ecovision
World Monograph Series. SPB Academic Publishing, Amsterdam.
Jude, D. J., R. H. Reider, and G. R. Smith. 1992. Establishment of Gobiidae in the
Great Lakes basin. Canadian Journal of Fisheries and Aquatic Sciences 49:416421.
Kazancheev, E. N. 1981. Fishes of the Caspian Sea: a key. [in Russian] Legkaya
Promyshlennost, Moscow.
King, R. B., J. M. Ray, and K. M. Stanford. 2006. Gorging on gobies: beneficial
effects of alien prey on a threatened vertebrate. Canadian Journal of Zoology
84:108-115.
Kornis, M. S., and M. J. Vander Zanden. 2010. Forecasting the distribution of the
invasive round goby (Neogobius melanostomus) in Wisconsin tributaries to Lake
Michigan. Canadian Journal of Fisheries and Aquatic Sciences 67:533-562.
Kovtun, I. F. 1979. Significance of the sex ratio in the spawning population of the
round goby, Neogobius melanostomus, in relation to year class strength in the
Sea of Azov. Journal of Ichthyology 19:161-163.
Kuhns, L. A., and M. B. Berg. 1999. Benthic invertebrate community responses to
round goby (Neogobius melanostomus) and zebra mussel (Dreissena
polymorpha) invasion in southern Lake Michigan. Journal of Great Lakes
Research 25:910-917.
Kulikova, N. I. 1985. Effect of chorionic gonadotropin on the growth and maturation
of oocytes in the Caspian round goby, Neogobius melanostomus (Gobiidae).
Journal of Ichthyology 25:86-98.
Kurunthachalam, K., L. Tao, E. Sinclair, S. D. Pastva, D. J. Jude, and J. P. Giesy.
2005. Perfluorinated compounds in aquatic organisms at various trophic levels in
a Great Lakes food chain. Archives of Environmental Contamination and
Toxicology 48:559-566.
Kwon, T. D., S. W. Fisher, G. W. Kim, H. Hwang, and K. Jang-Eok. 2006. Trophic
transfer and biotransformation of polychlorinated biphenyls in zebra mussel, round
goby and smallmouth bass in Lake Erie, USA. Environmental Toxicology and
Chemistry 25:1068-1079.
Lauer, T. E., P. J. Allen, et al. (2004). "Changes in mottled sculpin and johnny darter
trawl catches after the appearance of round gobies in the Indiana waters of Lake
Michigan." Transactions of the American Fisheries Society 133: 185-189.
Lederer, A. M., J. Janssen, T. Reed, and A. Wolf. 2008 Impacts of the introduced
round goby (Apollonia melanostoma) on dreissenids (Dreissena polymorpha and
Dreissena bugensis) and on macroinvertebrate community between 2003 and
2006 in the littoral zone of Green Bay, Lake Michigan. Journal of Great Lakes
Research 34:690-697.
Lederer, A., J. Massart, and J. Janssen. 2006 Impact of round gobies (Neogobius
melanostomus) on dreissenids (Dreissena polymorpha and Dreissena bugensis)
and the associated macroinvertebrate community across an invasion front.
Journal of Great Lakes Research 32:1-10.
http://infotrek.er.usgs.gov/wdnr_fishes/account.jsp?species_param=1490[7/10/2012 2:03:29 PM]
Fishes of Wisconsin
Lee, V. A., and T. B. Johnson. 2005. Development of a bioenergetics model for
round goby (Neogobius melanostomus). Journal of Great Lakes Research 31:125134.
Leslie, J. K., and C. A. Timmins. 2004. Description of age-0 round goby, Neogobius
melanostomus Pallas (Gobiidae), and ecotone utilization in St. Clair lowland
waters, Ontario. Canadian Field-Naturalist 118:318-325.
Li, W., M. J. Siefkes, A. P. Scott, and J. H. Teeter. 2003. Sex pheromone
communication in the sea lamprey: implications for integrated management.
Journal of Great Lakes Research 29(Supplement):85-94.
MacInnis, A. J., and L. D. Corkum. 2000a. Age and growth of round goby Neogobius
melanostomus in the upper Detroit River. Transactions of the American Fisheries
Society 129:852-858.
MacInnis, A. J., and L. D. Corkum. 2000b. Fecundity and reproductive season of the
round goby Neogobius melanostomus in the Upper Detroit River. Transactions of
the American Fisheries Society 129:136-144.
Maitland, P. S. 1977. The Hamlyn guide to freshwater fishes of Britain and Europe.
Hamlyn, London.
Meunier, B., S. Yavno, S. Ahmed, and L. D. Corkum. 2009. First documentation of
spawning and nest guarding in the laboratory by the invasive fish, the round goby
(Neogobius melanostomus). Journal of Great Lakes Research 35:608-612.
Miller, P. J. 1984. The tokology of gobioid fishes. Pages 119-153 in G. W. Potts and
R. J. Wooton, editors. Fish reproduction strategies and tactics. Academic Press,
London.
Miller, P. J. 1986. Gobiidae. Pages 1019-1095 in P. J. P. Whitehead, M. L. Bauchot,
J. C. Hureau, J. Nielsen, and E. Tortonese, editors. Fishes of the north-eastern
Atlantic and the Mediterranean. UNESCO, Paris.
Moiseyeva, Y. B., and V. I. Rudenko. 1996. The spawning of the round goby,
Gobius melanostomus, under aquarium conditions in winter. Journal of
Ichthyology 8:690-692.
Moskal'kova, K. I. 1996. Ecological and morphophysiological prerequisites to range
extension in the round goby Neogobius melanostomus under conditions of
anthropogenic pollution. Journal of Ichthyology 36:584-590.
Nichols, S. J., G. Kennedy, E. Crawford, J. Allen, J. R. P. French III, G. Black, M.
Blouin, J. Hickey, S. Chernyák, R. Haas, and M. Thomas. 2003. Assessment of
lake sturgeon (Acipenser fluvescens) spawning efforts in the Lower St. Clair River,
Michigan. Journal of Great Lakes Research 29:383-391.
Pallas, P. S. 1814 (1811-1814). Zoographia rosso-asiatica, sistens omnium
animalium in extenso Imperio Rossico et adjacentibus maribus observatorum
recensionem, domicilia, mores et descriptiones anatomen atque icones
plurimorum. Three volumes. Petropoli.
Pennuto, C. M., P. J. Krakowiak, and C. E. Janik. 2010. Seasonal abundance, diet,
and energy consumption of round gobies (Neogobius melanostomus) in Lake Erie
tributary streams. Ecology of Freshwater Fish 19:206-215.
Pinchuk, V. I. 1991. Species groupings in the genus Neogobius (Perciformes).
Journal of Ichthyology 31:1-15.
Ray, W. J., and L. D. Corkum. 1997. Predation of zebra mussels by round gobies,
Neogobius melanostomus. Environmental Biology of Fishes 50:267-273.
Ray, W. J., and L. D. Corkum. 2001. Habitat and site affinity of the round goby.
Journal of Great Lakes Research 27:329-334.
http://infotrek.er.usgs.gov/wdnr_fishes/account.jsp?species_param=1490[7/10/2012 2:03:29 PM]
Fishes of Wisconsin
Rollo, A., G. Andraso, J. Janssen, and D. Higgs. 2007. Attraction and localization of
round goby (Neogobius melanostomus) to conspecific calls. Behaviour 144:1-21.
Ross, R. M., J. H. Johnson, R. D. McCullough, and B. Edmonds. 2004. Diet
composition and fish consumption of double-crested cormorants from the Pigeon
and Snake Island colonies of eastern Lake Ontario in 2003. New York State
Department of Environmental Conservation, Special Report, New York.
Schaeffer, J. S., A. Bowen, M. Thomas, J. R. P. French III, and G. L. Curtis. 2005.
Invasion history, proliferation, and offshore diet of the round goby Neogobius
melanostomus in western Lake Huron, USA. Journal of Great Lakes Research
31:414-425.
Skazkina, E. P., and V. A Kostyuchenko. 1968. Food of N. melanostomus in the
Azov Sea. Voprosy Ikhtiologii 8:303-311.
Skora, K. E., and J. Stolarski. 1993. New fish species in the Gulf of Gdansk
Neogobius sp [(cf. Neogobius melanostomus (Pallas 1811)]. Morskiego Instytutu
Rybackiego Gdynia Bulletin of the Sea Fisheries Institute [Gydnia] 128:83.
Somers, C. M., M. N. Lozer, V. A. Kjoss, and J. S. Quinn. 2003. The invasive round
goby (Neogobius melanostomus) in the diet of nestling double-crested cormorants
(Phalacrocorax auritus) in Hamilton Harbour, Lake Ontario. Journal of Great
Lakes Research 29:392-399.
Spromberg, J. A., and W. J. Birge. 2005. Modeling the effects of chronic toxicity on
fish populations: the influence of life history strategies. Environmental Toxicology
and Chemistry 24:1532-1540.
Stammler, K. L., and L. D. Corkum. 2005. Assessment of fish size on shelter choice
and intraspecific interactions by round gobies Neogobius melanostomus.
Environmental Biology of Fishes 73:117-123.
Steingraeber, M. A., and P. A. Thiel. 2000. The round goby (Neogobius
melanostomus): another unwelcome invader in the Mississippi River Basin.
Transactions of the North American Wildlife and Natural Resources Conference
65:328-344.
Steingraeber, M., A. Runstrom, and P. Thiel. 1996. Round goby (Neogobius
melanostomus) distribution in the Illinois Waterway system of metropolitan
Chicago. U.S. Fish and Wildlife Publication, Fishery Resources Office, Onalaska,
Wisconsin.
Steinhart, G. B., E. A. Marschall, and R. A. Stein. 2004a. Round goby predation on
smallmouth bass offspring in nests during simulated catch and release angling.
Transactions of the American Fisheries Society 133:121-131.
Steinhart, G. B., R. A. Stein, and E. A. Marschall. 2004b. High growth rate of youngof-the-year smallmouth bass in Lake Erie: a result of round goby invasion?
Journal of Great Lakes Research 30:381-389.
Stepien, C. A., J. E. Brown, M. E. Neilson, and M. A. Tumeo. 2005. Genetic diversity
of invasive species in the Great Lakes versus their Eurasian source populations:
insights for risk analysis. Risk Analysis 25:1043-1060.
Stepien, C. A., and M. A. Tumeo. 2006. Invasion genetics of Ponto-Caspian gobies
in the Great Lakes: a 'cryptic' species, absence of founder effects, and
comparative risk analysis. Biological Invasions 8:61-78.
Stranai, I., and J. Andreji. 2004. The first report of round goby, Neogobius
melanostomus (Pisces, Gobiidae) in the waters of Slovakia. Folia Zoologica
53:335-338.
Svetovidov, A. N. 1964. Fishes of the Black Sea. [in Russian] Nauka Publications,
http://infotrek.er.usgs.gov/wdnr_fishes/account.jsp?species_param=1490[7/10/2012 2:03:29 PM]
Fishes of Wisconsin
Moscow.
Tsyplakov, E. P. 1974. Expansion of the ranges of some fishes in connection with
hydraulic engineering works on the Volga and acclimatization operations. Journal
of Ichthyology 14:343-351.
Vanderploeg, H. A., T. F. Nalepa, D. J. Jude, E. L. Mills, K. T. Holeck, J. R. Liebig, I.
A. Grigorovich, and H. Ojaveer. 2002. Dispersal and emerging ecological impacts
of Ponto-Caspian species in the Laurentian Great Lakes. Canadian Journal of
Fisheries and Aquatic Sciences 59:1209-1228.
Whelan, Gary E. 2007. Viral hemorrhagic septicemia (VHS) briefing paper. Michigan
Department of Natural Resources. Available:
http://www.michigan.gov/documents/dnr/Viral-Hemorrhagic-Septicemia-FactSheet-11-9-2006_178081_7.pdf.
Wickett, R. G., and L. D. Corkum. 1998. Nest defense by the non-indigenous fish,
the round goby, Neogobius melanostomus (Gobiidae), on a shipwreck in western
lake Erie. Canadian Field-Naturalist 112:653-656.
Young, J. A. M., J. R. Marentette, C. Gross, J. I. McDonald, A. Verma, S. E. MarshRollo, P. D. M. MacDonald, D. J. D. Earn, and S. Balshine. 2010. Demography
and substrate affinity of the round goby (Neogobius melanostomus) in Hamilton
Harbour. Journal of Great Lakes Research 36:115-122.
Yule, A. M., I. K. Barker, J. W. Austin, and R. D. Moccia. 2006. Toxicity of
Clostridium botulinum type E neurotoxin to Great Lakes fish: implications for avian
botulism. Journal of Wildlife Diseases 42:479-493.
Zielinski, B., W. Arbuckle, A. Belanger, L. Corkum, W. Li, and A. Scott. 2003.
Evidence for the release of sex pheromones by male round gobies (Neogobius
melanstomus). Fish Physiology and Biochemistry 28:237-239.
RETURN TO GOBY FAMILY
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