Natural Enemies of Zebra and Quagga Mussels: Predators, Parasites, and Ecological Competitors

Figures & data

Table 1. Fish documented in field studies in Europe (E) and North America (NA) as predators of planktonic Dreissena.

Species^a	Common names^b	Family	Where predation observed	References
Abramis bjoerkna (=Blicca bjoerkna)	Silver bream, white bream	Cyprinidae	E	Voronchuck et al. 1983
Abramis brama	Bream	Cyprinidae	E	Belyaev et al. 1970
Alburnus alburnus	Bleak	Cyprinidae	E	Voronchuck et al. 1983; Wilkońska and Strelnikova 2000
Alosa aestivalis	Blueback herring	Clupeidae	NA	Limburg and Ahrend 1994
Alosa pseudoharengus	Alewife	Clupeidae	NA	Creque and Czesny 2012; Eppehimer et al. 2019; Mills et al. 1995; Withers et al. 2015
Alosa sapidissima	American shad	Clupeidae	NA	Nack et al. 2015
Atherina boyeri	Sand smelt	Atherinidae	E	Chrisafi et al. 2007
Chondrostoma nasus	Nase	Cyprinidae	E	Mogilchenko 1986
Coregonus hoyi	Bloater	Salmonidae	NA	Eppehimer et al. 2019
Dorosoma cepedianum	Gizzard shad	Clupeidae	NA	Mills et al. 1995
Leuciscus idus	Ide, orfe	Cyprinidae	E	Mogilchenko 1986
Lota lota	Barbot	Gadidae	NA	George et al. 2013
Morone americana	White perch	Percichthyidae	NA	Limburg and Ahrend 1994; Limburg et al. 1997
Morone saxatilis	Striped bass	Percichthyidae	NA	Limburg et al. 1997
Notropis hudsonius	Spottail shiner	Cyprinidae	NA	Creque and Czesny 2012
Osmerus eperlanus	European smelt, sea smelt	Osmeridae	E	Wiktor 1958
Osmerus mordax	Rainbow smelt	Osmeridae	NA	Mills et al. 1995
Perca fluviatilis	Eurasian perch, European perch	Percidae	E	Wiktor 1958
Perca flavescens	Yellow perch	Percidae	NA	Creque and Czesny 2012; Marin Jarrin et al. 2015; Withers et al. 2015
Rutilus rutilus	Roach	Cyprinidae	E	Belyaev et al. 1970; Mogilchenko 1986; Wiktor 1958; Wilkońska and Strelnikova 2000; Zhdanova and Gusynskaya 1985
Scardinius erythrophthalmus	Rudd	Cyprinidae	E	Voronchuck et al. 1983; Wilkońska and Strelnikova 2000
Stizostedion lucioperca (=Lucioperca lucioperca)	Zander, pikeperch	Percidae	E	Draulans and Wouters 1988; Wiktor 1958

^aStandard scientific names follow Robins et al. (1991a, 1991b); names in parentheses are synonyms which appear in the corresponding references.

^bCommon names used in the accompanying references.

Table 2. Fish documented in field studies in Europe (E) or North America (NA) as predators of attached (i.e., juvenile and adult) Dreissena.

Species^a	Common names^b	Family	Where predation observed	References
Abramis bjoerkna (=Blicca bjoerkna)	Silver bream, white bream	Cyprinidae	E	Draulans and Wouters 1988; Filuk and Zmudzinski 1965; Gerasimov 2015; Karatayev et al. 1994; Kogan 1970; Mikheev 1963, 1977; Nagelkerke and Sibbing 1996; Nikitenko and Shcherbina 2016; Pliszka 1953; Prejs 1976; Shcherbina and Buckler 2006; Specziár et al. 1997; Zhokhov 2001
Abramis brama	Bream, common bream	Cyprinidae	E	Budzynska et al. 1956; Draulans and Wouters 1988; Filuk and Zmudzinski 1965; Gerasimov 2015; Gontya 1971; Mikheev 1963, 1977; Hartmann 1982; Karatayev et al. 1994; Martyniak et al. 1987; Nagelkerke and Sibbing 1996; Severenchuk and Kaftannikova 1983; Shcherbina and Buckler 2006; Specziár et al. 1997; Zheltenkova 1955; Zhokhov 2001
Acipenser brevirostrum	Shortnose sturgeon	Acipenseridae	NA	Gaygusuz et al. 2007; Bain, M. B., pers. comm.
Acipenser fulvescens	Lake sturgeon	Acipenseridae	NA	Bruestle et al. 2019; French 1993
Acipenser gueldenstaedti	Russian sturgeon	Acipenseridae	E	Yablonskaya 1985
Acipenser ruthenus	Sterlet	Acipenseridae	E	Karatayev et al. 1994; Egereva 1971; Mikheev 1977
Alburnus alburnus (=Alburnus lucidus)	Bleak	Cyprinidae	E	Sebestyén 1937
Ambloplites rupestris	Rock bass	Centrarchidae	NA	Watzin et al. 2008
Ameiurus nebulosus (=Ictalurus nebulosus)	Brown bullhead	Ictaluridae	NA	French 1993
Anguilla anguilla	European eel	Anguillidae	E	Biro 1974; de Nie 1982; Kublitskas 1959
Aplodinotus grunniens	Freshwater drum, drum	Sciaenidae	NA	Bartsch et al. 2005; French 1993; French and Bur 1993; Magoulick and Lewis 2002; Morrison et al. 1997; Shields and Beckman 2015; Thorp et al. 1998; Watzin et al. 2008
Babka (=Neogobius) gymnotrachelus	Racer goby	Gobiidae	E	Gaygusuz et al. 2007; Smirnov 1986
Benthophilus stellatus	Stellate tadpole-goby	Gobiidae	E	Kasyanov and Klevakin 2011; Smirnov 1986
Carassius carassius	Crucian carp	Cyprinidae	E	Naberezhny et al. 1971
Carassius auratus gibelio	Wild goldfish	Cyprinidae	E	Specziár et al. 1997
Carpoides carpio	River carpsucker	Catostomidae	NA	Thorp et al. 1998
Carpiodes cyprinus	Quillback carpsucker	Catostomidae	NA	Bartsch et al. 2005
Catostomus commersoni	White sucker	Catostomidae	NA	French 1993
Clupeonella cultriventris	Caspian sprat	Clupeidae	E	Sherstyuk and Severenchuk 1989
Coregonus clupeaformis	Lake whitefish	Salmonidae	NA	French 1993; Owens and Dittman 2003; Pothoven and Nalepa 2006; Pothoven et al. 2001; Lumb et al. 2007; Rennie et al. 2012
Coregonus lavaretus	Powan	Salmonidae	E	Peczalska 1961; Pliszka 1953
Coregonus macrophthalmus	Gangfisch, benthivorous whitefish	Salmonidae	E	Baer et al. 2022
Cyprinus carpio	Common carp	Cyprinidae	E and NA	Aleksenko 2004; Bartsch et al. 2005; Botnariuc et al. 1964; Marsden 1997; Mikheev 1963; Sebestyén 1937; Specziár et al. 1997; Stein et al. 1975; Thorp et al. 1998; Tucker et al. 1996
Huso huso	Beluga	Acipenseridae	E	Yablonskaya 1985
Ictiobus bubalus	Smallmouth buffalo	Catostomidae	NA	Thorp et al. 1998
Ictalurus furcatus	Blue catfish	Ictaluridae	NA	Herod et al. 1997; Magoulick and Lewis 2002
Ictalurus punctatus	Channel catfish	Ictaluridae	NA	Thorp et al. 1998
Lepomis auritus	Redbreast sunfish	Centrarchidae	NA	Boles and Lipcius 1997; Schmidt, R., pers. comm.
Lepomis gibbosus	Pumpkinseed	Centrarchidae	NA and E	Andraso 2005; Berchtold et al. 2015; Boles and Lipcius 1997; Colborne et al. 2015; Locke et al. 2014; Naddafi and Rudstam 2014c^a; Rezsu and Specziár 2006; Spataru 1967; Watzin et al. 2008
Lepomis macrochirus	Bluegill	Centrarchidae	NA	Andraso 2005; Bartsch et al. 2005; Magoulick and Lewis 2002; Mercer et al. 1999; Thorp et al. 1998
Lepomis microlophus	Redear sunfish	Centrarchidae	NA	Magoulick and Lewis 2002
Leuciscus idus	Ide, orfe	Cyprinidae	E	Mikheev 1963, 1977; Pliszka 1953; Shcherbina and Buckler 2006; Zhokhov 2001
Leuciscus leuciscus	Dace	Cyprinidae	E	Baer et al. 2022b)
Macrhybopsis storeriana	Silver chub	Cyprinidae	NA	Kočovský 2019
Morone americana	White perch	Percichthyidae	NA	French 1993
Morone chrysops	White bass	Percichthyidae	NA	French 1993
Moxostoma sp.	Redhorse sucker	Catostomidae	NA	Bartsch et al. 2005; Thorp et al. 1998
Moxostoma valenciennesi	Greater redhorse	Catostomidae	NA	French 1993
Mylopharyngodon piceus	Black carp, black amur, Chinese roach	Cyprinidae	E and NA	Evtushenko et al. 1994; Konradt and Mukhammedova 1974; Krepis et al. 1981; Nebolsina et al. 1991; Poulton et al. 2019
Neogobius fluviatilis	Monkey goby	Gobiidae	E	Smirnov 1986; Yablonskaya 1985
Neogobius kessleri	Bighead goby	Gobiidae	E	Gavlena 1977; Yablonskaya 1985
Neogobius melanostomus	Round goby	Gobiidae	E and NA	Bunnell et al. 2015; Campbell et al. 2009; Camp et al. 1999; Djuricich and Janssen 2001; Jude et al. 1995; Kuhns and Berg 1999; Lederer et al. 2006, 2008; Miano et al. 2021; Muzzall et al. 1995; Naddafi and Rudstam 2014b, 2014c^a; Rakauskas et al. 2008; Ray and Corkum 1997; Rybczyk et al. 2020; Shemonaev and Kirilenko 2009; Smirnov 1986; Turschak and Bootsma 2015; Yablonskaya 1985
Perca flavescens	Yellow perch	Percidae	NA	French 1993; Morrison et al. 1997; Watzin et al. 2008
Perca fluviatilis	Eurasian perch, European perch	Percidae	E	Naberezhny et al. 1971; Sebestyén 1937
Percina caprodes	Logperch		NA	Burkett and Jude 2015
Platichthys flesus (=Pleuronectes flesus)	European flounder, flounder	Pleuronectidae	E	Filuk and Zmudzinski 1965
Proterorhinus marmoratus (=Priolepis marmoratus)	Tubenose goby	Gobiidae	E	Yablonskaya 1985
Pylodictis olivaris	Flathead catfish	Ictaluridae	NA	Bartsch et al. 2005; Thorp et al. 1998
Rhodeus sericeus	Bitterling	Cyprinidae	E	Naberezhny et al. 1971
Rutilus rubilio	Adriatic roach	Cyprinidae	E	Daoulas and Economidis 1984
Rutilus rutilus	Roach	Cyprinidae	E	Aleksenko 2004; Baer et al. 2022; Balkuvienė and Pernaravičiūté 1994; Dmitrenko 1967; Draulans and Wouters 1988; Filuk and Zmudzinski 1965; Gerasimov 2015; Grigorash 1963; Karatayev et al. 1994; Kharchenko 1975; Kodukhova and Karabanov 2017; Mikheev 1963; Nagelkerke and Sibbing 1996; Nikitenko and Shcherbina 2016; Olszewski 1978; Pliszka 1953; Pirozhnikov 1971; Prejs 1976; Rask 1989; Specziár et al. 1997; Shcherbina and Buckler 2006; Stanczykowska 1987; Zheltenkova 1949; Zhokhov 2001
Stizostedion vitreum	Walleye	Percidae	NA	French 1993
Tinca tinca	Tench	Cyprinidae	E	Baer et al. 2022; Pliszka 1953; Severenchuk and Kaftannikova 1983
Vimba elongata	No common name	Cyprinidae	E	Ritterbusch-Nauwerck 1991
Vimba vimba	Vimba	Cyprinidae	E	Aleksenko 2004; Kublitskas 1959; Okgerman et al. 2013
Fish that consume dreissenids only occasionally
Acipenser sturio	European Atlantic sturgeon	Acipenseridae	NA	Ferreira-Rodriguez et al. 2016
Gymnocephalus cernua	Ruffe	Percidae	E	Baer et al. 2022b
Esox lucius	Northern pike	Esocidae	E	Millane et al. 2012
Notropis hudsonius	Spottail shiner	Cyprinidae	NA	Creque and Czesny 2012; Turschak and Bootsma 2015
Prosopium cylindraceum	Round whitefish	Salmonidae	NA	Turschak and Bootsma 2015
Salmo trutta	Brown trout	Salmonidae	E	Millane et al. 2012
Salvelinus profundus	Deep-water Arctic charr	Salmonidae	E	Baer et al. 2022b
Salvelinus umbla	Normal Arctic charr	Salmonidae	E	Baer et al. 2022b
Squalius cephalus	Chub	Cyprinidae	E	Baer et al. 2022b

^aStandard scientific names follow Robins et al. (1991a, 1991b); names in parentheses are synonyms which appear in the accompanying references.

^bCommon names used in the corresponding references.

Table 3. Relationship between predation on attached (i.e., juvenile and adult) Dreissena and fish length and/or age in field populations.

Species^a	Fish length and/or age when start feeding on Dreissena	Location	References
Abramis bjoerkna (=Blicca bjoerkna)	>70 mm	Mikolajskie Lake, Poland	Prejs 1976
	>105 mm (2+ years)	Rybinsk Reservoir, Russia	Zhokhov 2001
Abramis brama	18% of ≥4-year old (<1% of diet)	Goplo Lake, Poland	Budzynska et al. 1956
	>370 mm	Pierzchaly Reservoir, Poland	Martyniak et al. 1987
	≥315 mm long	Rybinsk Reservoir, Russia	Zhokhov 2001
Aplodinotus grunniens	Mainly >250 mm but consumption increases with fish length: dreissenids present in 16% of 200–249 mm long fishes; 250–374 mm individuals feed mainly on dreissenids; >375 mm fish feed almost exclusively on dreissenids (97–99% of the total dry weight of food ingested). The smallest ind. capable of crushing dreissenid shells is 265 mm.	Lake Erie, USA	French and Bur 1993
	>250 mm	Lake Erie, USA	Morrison et al. 1997
	>200 mm; ∼50% of large fish consumed mussels.	Lake Dardanelle, USA	Magoulick and Lewis 2002
Coregonus clupeaformis	73–149 mm (age-0 fish) mainly consume large-bodied cladoceran zooplankton; ≤350 mm feed on soft-bodied macroinvertebrates and a few molluscs; >350 mm mainly eat molluscs, particularly D. r. bugensis	Lake Huron, USA	Pothoven and Nalepa 2006
Cyprinus carpio	No conclusive evidence that 381–666 mm fish consumed more dreissenids than 299–373 mm	Mississippi River, USA	Tucker et al. 1996
Ictalurus furcatus	22% of 25–50 cm feed on dreissenids	Ohio River, USA	Herod et al. 1997
Leuciscus idus	≥220 mm	Rybinsk Reservoir, Russia	Zhokhov 2001
Mylopharyngodon piceus	Age 2	Tsimlyanskoe Reservoir, Russia	Mukhammedova et al. 1989
Neogobius melanostomus	47–65 mm fish feed on dreissenids and of soft-bodied invertebrates; in 80–90 mm dreissenids comprise up to 82% of diet >100 mm can feed almost exclusively on dreissenids.	Great Lakes, USA	Jude et al. 1995
	Age 2+ diet was dominated by zebra mussels (75% by weight)	Curonian Lagoon, Lithuania	Rakauskas et al. 2008
Perca flavescens	>150 mm	Lake Erie, USA	Morrison et al. 1997
Rutilus rubilio	<159 mm fish feed on plant and animal food, the largest ones mainly consuming dreissenids	Lake Trichonis, Greece	Daoulas and Economidis 1984
Rutilus rutilus	≥100 mm 7% of Dreissena in guts of 200 mm roach, 72% in 250 mm, and 99% in 300 mm roach	Dnieper-Bug estuary region, Ukraine	Aleksenko 2004
	57% of 4-year old and 100% of ≥8-year-old roach had dreissenids in their guts (54–97% of the gut contents)	Goplo Lake, Poland	Budzynska et al. 1956
	140–160 mm	Lake Harsz, Poland	Budzynska et al. 1956
	140–160 mm	Goplo Lake, Poland	Pliszka 1953
	140–160 mm	Mikolajskie Lake, Poland	Prejs 1976
	180–320 mm roach feed almost exclusively on dreissenids	Lake Harsz, Poland	Pliszka 1953
	>160 mm feed on molluscs (95% dreissenids)	Mikolajskie Lake, Poland	Prejs 1976
	<160 mm rarely fed on dreissenids; <180 mm fed on plants; 160–179 mm had 6% dreissenids in diet, 230–240 mm—75%, >280 mm—95%	Lake Sniardwy, Poland	Prejs et al. 1990
	Start feeding at >150 mm; feed almost exclusively when >200 mm	Uchinskoe Reservoir, Russia	Spanowskaya 1963
	160–179 mm roach had 6% dreissenids in the diet; 280–350 mm had 97%	Lakes in Poland	Stanczykowska 1987
	>140 mm long (at 2+ years)	Rybinsk Reservoir, Russia	Zhokhov 2001

^aStandard scientific names following Robins et al. (1991a, 1991b); names in parentheses are synonyms which appear in the accompanying references.

Table 4. Seasonal changes in fish predation on attached (i.e., juvenile and adult) Dreissena: a reflection of preferred prey seasonal scarcity?

Species	Food spectrum and when feed on Dreissena	Location	References
Abramis brama	Food spectrum was very wide with chironomid larvae as preferred prey year long; Dreissena was the most important mollusc in the diet and was eaten primarily May through September	Pierzchaly Reservoir, Poland	Martyniak et al. 1987
Anguilla anguilla	Exhibit stronger preference for fish; bivalves like Dreissena contributed significantly to the diet in September and October	Tjeukemeer, Netherlands	de Nie 1982
Aplodinotus grunniens	250–374 mm drum feed at reef sites predominately on dipterans in May and dreissenids in July and September, whereas in locations with silt or sand shifted from dreissenids in July to small fish in September	Lake Erie, USA	French and Bur 1993
	The lowest dreissenid predation was in October (of studied May, July and October)	Lake Erie, USA	Morrison et al. 1997
Cyprinus carpio	Dreissenid predation was higher in summer months (in this July-January study)	Kuibyshev Reservoir, Russia	Mikheev 1963
	Dreissenids were observed in stomachs in summer and December	Lake Michigan, USA	Marsden 1997
Ictalurus furcatus	Feed on abundant zebra mussels during summer and shifted to shad Dorosoma spp. during winter	Lake Dardanelle, USA	Magoulick and Lewis 2002
Neogobius melanostomus	Dreissena spp. and Bythotrephes longimanus dominated the diet in the summer	Lake Huron, USA	Pothoven and Madenjian 2013
	Large gobies consumed relatively high proportions of dreissenids in all seasons; small gobies mostly feed on dreissenids in spring and autumn	Lake Michigan, USA	Bunnell et al. 2015
Perca flavescens	The highest dreissenid predation was in May (stomachs examined in May, July and October)	Lake Erie, USA	Morrison et al. 1997
Rutilus rubilio	Older fish consumed molluscs (dreissenids and gastropods) all year long in large quantities; small and medium fish consumed molluscs only from October to February.	Lake Trichonis, Greece	Daoulas and Economidis 1984
Rutilus rutilus	Dreissenids are the main food in July when the growth rates are high	Goplo Lake, Poland	Budzynska et al. 1956
	Feed on dreissenids primarily in autumn and winter	Firth of Szczecin, Poland	Wiktor 1969

Table 5. Size of Dreissena consumed by fish.

Species^a	Dreissena length	Where predation observed	References
Abramis bjoerkna (=Blicca bjoerkna)	2–14 mm	Kuibyshev Reservoir, Russia	Mikheev 1977
	Largest dreissenids ingested by 128–246 mm fish were, respectively, 4.1 and 9.7 mm in width	Lab study, the Netherlands	Nagelkerke and Sibbing 1996
	4–14 mm	Upper Volga Basin, Russia	Shcherbina and Buckler 2006
Abramis brama	6–8 mm (max 13 mm)	Kuibyshev Reservoir, Russia	Mikheev 1977
	2–10 mm	Upper Volga Basin, Russia	Shcherbina and Buckler 2006
	Largest dreissenids ingested by 155–298 mm fish were, respectively, 3.0 and 7.0 mm in width	Lab study, Netherlands	Nagelkerke and Sibbing 1996
Acipenser ruthenus	2–6 mm (max 15 mm)	Kuibyshev Reservoir, Russia	Mikheev 1977
Aplodinotus grunniens	Maximum 21.4 mm	Lake Erie, USA	French and Love 1995
	Drum <350 mm total length selected 4–6 mm dreissenids; larger fish were less selective, consuming sizes in proportion to availability	Lake Erie, USA	Morrison et al. 1997
Cyprinus carpio	1–6 mm (young carp)	Kuibyshev Reservoir, Russia	Mikheev 1963
	Large carp (∼54 cm) fed on large dreissenids (28 mm)	Dnieper-Bug estuary region, Ukraine	Aleksenko 2004
	Large carp (299–666 mm) fed on 1.5 to 42.5 mm dreissenids (average 11.8 mm)	Mississippi River, USA	Tucker et al. 1996
Cyprinus carpio, Moxostoma sp., Aplodinotus grunniens, and possibly other species	>12 mm (enclosure experiments in low-density mussel populations)	Mississippi River, USA	Thorp et al. 1998
Ictalurus furcatus	25–50 cm fish fed on dreissenids 3–15 mm (preference for 3–7 mm)	Ohio River, USA	Herod et al. 1997
Ictiobus bubalus, Ictalurus punctatus, Aplodinotus grunniens, Carpoides carpio, Pylodictis olivaris, and possibly other species	Enclosure experiments in high-density mussel populations did not provide evidence of size-selective predation	Ohio River, USA	Thorp et al. 1998
Lepomis microlophus	7 mm long juvenile fish consumed dreissenids <3 mm long; 180–260 mm long adults fed on dreissenids up to 20 mm	Lab study, USA	French and Morgan 1995
Lepomis gibosus	140–185 mm fish consume 5–13 mm long D. polymorpha	Lab study, USA	Naddafi and Rudstam 2014c
Leuciscus idus	8–15 mm (max 17 mm)	Kuibyshev Reservoir, Russia	Mikheev 1977
	10–30 mm	Upper Volga Basin, Russia	Shcherbina and Buckler 2006
Neogobius melanostomus	60–100 mm gobies consumed 4.5–12.5 mm dreissenids	Lab study, USA	Ghedotti et al. 1995
	55–103 mm gobies ate dreissenids <10.0 mm long; 85–103 mm gobies ate 10.0–12.9 mm dreissenids, but preferred smaller sizes	Lab study, Canada	Ray and Corkum 1997
	Gobies 94–135 mm ate 5–13 mm D. polymorpha	Lab study, USA	Naddafi and Rudstam 2014c
	0.5–14.3 mm (mean 3.4 mm)	Saginaw Bay of Lake Huron, USA	Foley et al. 2017
Perca flavescens	<200 mm fish consumed dreissenids <6 mm larger fish preferred 1–15 mm mussels	Lake Erie, USA	Morrison et al. 1997
Rutilus rutilus	6–18 mm (max 19 mm)	Kuibyshev Reservoir, Russia	Mikheev 1977
	180–209 mm and ≥210 mm roach preferred 10–11 mm and 11–17 mm dreissenids, respectively.	Lake Sniardwy, Poland	Prejs et al. 1990
	5–20 mm	Upper Volga Basin, Russia	Shcherbina and Buckler 2006
	Largest dreissenids ingested by fish from 146 to 235 mm were, respectively, 5.4 and 10.3 mm in width	Lab study, Netherlands	Nagelkerke and Sibbing 1996
	Up to 24 mm	Dnieper-Bug estuary region, Ukraine	Aleksenko 2004

^aStandard scientific names following Robins et al. (1991a, 1991b); names in parentheses are synonyms which appear in the accompanying references.

Table 6. Impact of fish predation on Dreissena populations.

Species	Notes	Where predation observed	References
Aplodinotus grunniens	Long-term suppression of dreissenid populations by freshwater drum not likely; drum >250 mm long can consume large numbers of dreissenids, but represented only 19% of population; most efficient predators, drum >350 mm long, were not common.	Lake Erie, USA	French and Bur 1993
Aplodinotus grunniens (and possibly other fish)	Predation affected D. r. bugensis (particularly <10 mm long) but not D. polymorpha abundance.	Lake Erie, USA	Mitchell et al. 2000
Aplodinotus grunniens, Ictalurus furcatus, Lepomis microlophus (and possibly other fish)	Densities of large (>5 mm) zebra mussels were significantly lower in in predator exposure treatments than in predator exclosure treatments during 2 years of trials	Lake Dardanelle, USA	Magoulick and Lewis 2002
Cyprinus carpio, Moxostoma sp., Aplodinotus grunniens (and possibly other fish)	Fish-exclusion cages had an overall significantly higher densities and biomass of dreissenids than fish-accessible cages	Mississippi River, USA	Thorp et al. 1998
Ictiobus bubalus, Ictalurus punctatus, Aplodinotus grunniens, Carpoides carpio, Pylodictis olivaris, and possibly other species	Fish-exclusion cages had an overall significantly higher densities and biomass of dreissenids than fish-accessible cages	Ohio River, USA	Thorp et al. 1998
Lepomis auritus, Lepomis gibbosus and possibly other fish species	14% reduction of dreissenids likely due to predation (two weeks open cage field studies in summer); current levels of finfish unlikely to limit dreissenid populations	Hudson River, USA	Boles and Lipcius 1997
Mylopharyngodon piceus	4-year-old fish consumed 1.4 kg/day of Dreissena in experimental ponds and 1.8 kg/day in hatcheries	Artificial fish rearing areas, Moldavia	Krepis et al. 1981
Neogobius melanostomus	Preference for small dreissenids has the potential to alter the size structure of dreissenid populations in specific locales	Lab study, Canada	Ray and Corkum 1997
	During 2001–2004 densities of D. r. bugensis decreased 94% in the nearshore rocky zone	Easter basin of Lake Erie, USA	Barton et al. 2005
	Increase in goby abundance was associated with the decline of dreissenids biomass from 344–524 to 34–73 g shell-on dry weight/m^2 from 2009–2011 to 2016–2018	Onondaga Lake, USA	Rudstam and Gandino 2020
	Have little (if any) lake-wide effect at the scale of deep Great Lakes	Great Lakes, USA	Bunnell et al. 2005; Foley et al. 2017; Johnson et al. 2005b; Karatayev et al. 2022a
Rutilus rutilus	Benthophagous fish (mainly roach) eat 80% of the annual production of <15 mm dreissenids	Uchinskoe Reservoir, Russia	Lvova 1977
Rutilus rutilus, Anguilla anguilla, and others	Fish consumed 1–2 × 10^12J of lake annual dreissenid production (total 41–102 × 10^12J)	Lake IJsselmeer, the Netherlands	Smit et al. 1993
Species not mentioned	130 × 10^6 kg of dreissenids (ca. 90% of the annual production) are eaten by fish yearly	Caspian Sea, Russia	Yablonskaya 1985

Table 7. Birds documented to eat Dreissena in the field and where predation was observed (E—Europe, NA—North America).

Species	Common name	Duck type	Where predation observed	References
Anas platyrhynchos	Mallard	Puddle duck	E	Jacoby and Leuzinger 1972; Kozulin 1995; Zuur et al. 1983
Anas rubripes	Black duck	Puddle duck	NA	Petrie and Knapton 1999
Anas strepera	Gadwall	Puddle duck	E	Jacoby and Leuzinger 1972
Aythya affinis	Lesser scaup	Diving duck	NA	Badzinski and Petrie 2006; Custer and Custer 1996; Hamilton and Ankney 1994; Hamilton et al. 1994; Mazak et al. 1997; Mitchell and Carlson 1993; Petrie and Knapton 1999; Wormington and Leach 1992
Aythya americana	Redhead	Diving duck	NA	Custer and Custer 1996; Petrie and Knapton 1999
Aythya collaris	Ring-necked duck	Diving duck	NA	Wormington and Leach 1992
Aythya ferina	Pochard, European pochard	Diving duck	E	bij de Vaate 1991; Cleven and Frenzel 1992; Géroudet 1978; Jacoby and Leuzinger 1972; Leuzinger and Schuster 1970; Pedroli 1981a; Smit et al. 1993; Suter 1982a, 1982b; Zuur et al. 1983; van Eerden and de Leeuw 2010; van Eerden et al. 1997; Wiktor 1969
Aythya fuligula	Tufted duck	Diving duck	E	Burla and Lubini-Ferlin 1976; Cleven and Frenzel 1992; de Leeuw and Renema 1997; Draulans and De Bont 1980; Géroudet 1966, 1978; Jacoby and Leuzinger 1972; Leuzinger and Schuster 1970; Olney 1963; Pedroli 1981a; Smit et al. 1993; Suter 1982a, 1982b; van Eerden and de Leeuw 2010; van Eerden et al. 1997; Zuur et al. 1983
Aythya marila	Greater scaup	Diving duck	E and NA	Badzinski and Petrie 2006; bij de Vaate 1991; Custer and Custer 1996; Géroudet 1966, 1978; Hamilton and Ankney 1994; Hamilton et al. 1994; Jacoby and Leuzinger 1972; Leuzinger and Schuster 1970; Marchowski et al. 2015; Mazak et al. 1997; Mitchell and Carlson 1993; Pedroli 1981a; Petrie and Knapton 1999; Smit et al. 1993; Suter 1982a; van Eerden and de Leeuw 2010; van Eerden et al. 1997; Wormington and Leach 1992
Aythya nyroca	Ferruginous duck, white-eye	Diving duck	E	Jacoby and Leuzinger 1972; Suter 1982a
Aythya valisineria	Canvasback	Diving duck	NA	Custer and Custer 1996
Bucephala albeola	Bufflehead	Diving duck	NA	Custer and Custer 1996; Hamilton and Ankney 1994; Hamilton et al. 1994; Mazak et al. 1997; Petrie and Knapton 1999; Wormington and Leach 1992
Bucephala clangula	Goldeneye, common goldeneye	Diving duck	E and NA	bij de Vaate 1991; Custer and Custer 1996; de Leeuw and Renema 1997; Géroudet 1966, 1978; Jacoby and Leuzinger 1972; Hamilton and Ankney 1994; Leuzinger and Schuster 1970; Smit et al. 1993; Suter 1982a, 1982b; van Eerden and de Leeuw 2010; van Eerden et al. 1997; Wormington and Leach 1992
Calidris alpina	Dunlin	Shore bird	NA	Wormington and Leach 1992
Calidris maritima	Purple sandpiper	Shore bird	NA	Wormington and Leach 1992
Charadrius vociferus	Killdeer	Shore bird	NA	Wormington and Leach 1992
Clangula hyemalis	Oldsquaw, longtailed duck	Diving duck	E and NA	Essian et al. 2016; Hamilton and Ankney 1994; Knapton R. W., pers. comm.; Suter 1982a
Cygnus olor	Mute swan	Swan	E	Jacoby and Leuzinger 1972; von Wicht 1972; Włodarczyk and Janiszewski 2014
Euphagus carolinus	Rusty blackbird	Passeriform	NA	Wormington and Leach 1992
Fulica americana	American coot	Diving rail	NA	Wormington and Leach 1992
Fulica atra	Coot, baldcoot	Diving rail	E	Borowiec 1975; de Leeuw and Renema 1997; Géroudet 1966; Jacoby and Leuzinger 1972; Königstein 1986; Kornobis 1977; Krauß 1979; Leuzinger and Schuster 1970; Piesik 1983; Smit et al. 1993; Stempniewicz 1974; Suter 1982a, 1982b; van Eerden and de Leeuw 2010; van Eerden et al. 1997; Zuur et al. 1983
Gallinula chloropus	Common gallinule, moorhen	Diving rail	E	Jacoby and Leuzinger 1972
Larus argentatus	Herring gull	Gull	E and NA	Jacoby and Leuzinger 1972; Marchowski et al. 2015; Wormington and Leach 1992
Larus canus	Common gull, mew gull	Gull	E	Jacoby and Leuzinger 1972; Marchowski et al. 2015
Larus delawarensis	Ring-billed gull	Gull	NA	Wormington and Leach 1992
Larus ridibundus	Blackheaded gull	Gull	E	Jacoby 2005
Larus spp.	Gulls	Gull	E	de Leeuw and Renema 1997; Marchowski et al. 2015
Lophodytes cucullatus	Hooded merganser	Diving bird	NA	Wormington and Leach 1992
Melanitta fusca	White-winged scoter, velvet scoter	Diving duck	E and NA	Géroudet 1966; Hamilton and Ankney 1994; Knapton R. W., pers. comm.; Wormington and Leach 1992
Melanitta nigra	Black scoter, common scoter	Diving duck	NA	Wormington and Leach 1992
Melanitta perspicillata	Surf scoter	Diving duck	NA	Essian et al. 2016; Guillemette et al. 1994; Wormington and Leach 1992
Mergus merganser	Common merganser, goosander	Diving bird	E	Jacoby and Leuzinger 1972
Mergus serrator	Red-breasted merganser, sawbill	Diving bird	E	Jacoby and Leuzinger 1972
Netta rufina	Red-crested pochard, scarp duck	Diving duck	E	Jacoby and Leuzinger 1972; Leuzinger and Schuster 1970; Suter 1982a
Phalacrocorax auritus	Double-crested cormorants	Diving bird	NA	Essian et al. 2016
Podiceps ruficollis	Grebe	Diving bird	E	Jacoby and Leuzinger 1972
Somateria mollissima	Eider, common eider	Diving duck	E	Géroudet 1966, 1978; Jacoby and Leuzinger 1972; Leuzinger and Schuster 1970
Sturnus vulgaris	European starling	Passeriform	NA	Wormington and Leach 1992
Tadorna tadorna	Shelduck, common shelduck	Puddle duck	E	Jacoby and Leuzinger 1972

Table 8. Size of Dreissena consumed by birds.

Species	Common name	Length of mussels	Where predation observed	References
Aythya affinis	Lesser scaup	≤24 mm (7–15 mm most common)	Lake Erie, Canada	Mazak et al. 1997
		9.0 mm (spring), 6.5 mm (autumn) 10.9 mm (Lake Erie) 6.9 mm (Lake Ontario), 5.5 mm (Lake St. Clair)	Great Lakes, Canada	Badzinski and Petrie 2006
		Mean 12 mm at Long Point and 14 mm at Point Pelee	Lake Erie, Canada	Hamilton and Ankney 1994
		Mean 8 mm	Lake St. Clair, Canada	Hamilton and Ankney 1994
Aythya ferina	Pochard, European pochard	≤27 mm, but tended to take <15 mm, especially at depths of <2.5 m	Lake IJsselmeer area, the Netherlands	van Eerden et al. 1997
Aythya fuligula	Tufted duck	10–20 mm preferred	Lake Zurich, Switzerland	Burla and Lubini-Ferlin 1976
		11–25 mm	Waterbodies in England and Northern Ireland	Olney 1963
		Males: ≤25 mm (15–20 mm preferred) Females: ≤2.5 mm (10–17 mm preferred)	Experimental chambers	Draulans 1982
		≤30 mm (slight preference for 7–16 mm)	Experimental chambers	de Leeuw 1999
		≤27 mm (<15 mm preferred, especially at depths of <2.5 m)	Lake IJsselmeer area, the Netherlands	van Eerden et al. 1997
Aythya marila	Greater scaup	≤27 mm (10–21 mm most common)	Lake Erie, Canada	Mazak et al. 1997
		≤30 mm (slight preference for 7–16 mm)	Experimental chambers	de Leeuw 1999
		≤27 mm (preferred <15 mm, especially at depths of <2.5 m)	Lake IJsselmeer area, the Netherlands	van Eerden et al. 1997
		Mean 12 mm at Long Point	Lake Erie, Canada	Hamilton and Ankney 1994
		Mean 10 mm	Lake St. Clair, Canada	Hamilton and Ankney 1994
Bucephala albeola	Bufflehead	≤21 mm (1–9 mm most common)	Lake Erie, Canada	Mazak et al. 1997
		Mean 9 mm at Long Point and 10.5 mm at Point Pelee	Lake Erie, Canada	Hamilton and Ankney 1994
Bucephala clangula	Goldeneye	≤27 mm (preferred <15 mm, especially at depths of <2.5 m)	Lake IJsselmeer area, the Netherlands	van Eerden et al. 1997
		Mean 14 mm at Long Point and 11 mm at Point Pelee	Lake Erie, Canada	Hamilton and Ankney 1994
Clangula hyemalis	Oldsquaw	Mean 14 mm at Port Stanley	Lake Erie, Canada	Hamilton and Ankney 1994
Melanitta fusca	White-winged scoter	Mean 13.5 mm at Port Stanley	Lake Erie, Canada	Hamilton and Ankney 1994
Aythya affinis, Aythya marila, Bucephala albeola	Lesser scaup, greater scaup, bufflehead	11–21 mm (11–13 mm preferred)	Lake Erie, Canada	Hamilton et al. 1994
Aythya affinis, Aythya marila, Aythya americana, Bucephala albeola	Lesser scaup, greater scaup, bufflehead, redhead	<15 mm	Lake Erie, Canada	Petrie and Knapton 1999
Aythya affinis, Aythya marila, Bucephala albeola, Bucephala clangula	Lesser scaup, greater scaup, bufflehead, goldeneye	Mean 8–12 mm for individuals specifically foraging on dreissenids	Lake Erie and Lake St. Clair, USA/Canada	Custer and Custer 1996

Figure 1. Blue crab Callinectes sapidus eating D. polymorpha (Credit: D. P. Molloy).

Table 9. Crayfish species documented eating Dreissena.

Species	Where predation observed	References
Astacus astacus	Germany	Chucholl and Chucholl 2021^a
Astacus leptodactylus	Ukraine	Grinbart and Suprunovitch 1981
	Kazakhstan	Malinowskaya 1976
	Hungary	Sebestyén 1937
Austropotamobius pallipes	Ireland	Reynolds and Donohoe 2001^a
Cambarus affinis	Germany	Pieplow 1938
Cambarus robustus	United States	Hazlett 1994^a
Faxonius immunis	Germany	Chucholl and Chucholl 2021^a
Faxonius limosus	Germany	Chucholl and Chucholl 2021^a; Linzmaier and Jeschke 2020^a
Orconectes limosus	Poland	Kornobis 1977; Piesik 1974^a, 1983; Szlauer 1974
	Netherlands	Smit et al. 1993
Orconectes propinquus	Canada United States	MacIsaac 1994^a; Martin and Corkum 1994^a; Perry et al. 1997^a
Orconectes rusticus	United States	Green et al. 2008; Naddafi and Rudstam 2014c^a; Perry et al. 1997, 2000
Orconectes virilis	United States	Hazlett 1994^a; Love and Savino 1993^a; Perry et al. 1997^a
Pacifastacus leniusculus	SwedenGermanySwedenU.K.	Chucholl and Chucholl 2021^a; Schreiber et al. 1998^a; zu Ermgassen and Aldridge 2011^a
Procambarus clarkii	Italy	Chucholl 2013; Gonçalves et al. 2016^a, 2017^a
Procambarus virginalis	Germany	Chucholl and Chucholl 2021^a; Linzmaier and Jeschke 2020^a

^aLab observations only.

Table 10. Endosymbionts including commensal (C) and parasitic (P) reported from attached Dreissena polymorpha and D. r. bugensis populations in Europe.

Endosymbionts	Type of symbiont	D. polymorpha	D. r. bugensis
Ciliates
Conchophthirus acuminatus	C	Switzerland (Claparède and Lachmann 1858)	Ukraine (Karatayev et al. 2000b; Yuryshynets 2019)
		Bulgaria (Raabe 1934)	Russia (Pryanichnikova et al. 2011; Tyutin et al. 2013a)
		Poland (Dobrzanska 1958; Raabe 1934, 1956)
		Hungary (Raabe 1950)
		Denmark (Fenchel 1965)
		Belarus (Burlakova et al. 1998; Karatayev et al. 2000a, 2000b, 2003a, 2007; Mastitsky 2012)
		Russia (Laruelle et al. 1999; Pryanichnikova et al. 2011; Tyutin et al. 2013a)
		Ukraine (Karatayev et al. 2000a; Yuryshynets et al. 2008)
		Ireland (Burlakova et al. 2006a; Conn et al. 2008)
		Sweden (Mastitsky et al. 2008)
		Lithuania (Chuševė et al. 2012)
		France (Minguez et al. 2009, 2011, 2013)
		Germany (Molloy et al. unpublished data)
		Italy (Molloy et al. unpublished data)
		Greece (Molloy et al. unpublished data)
		The Netherlands (Molloy et al. unpublished data)
Hypocomagalma dreissenae	P	Hungary (Raabe 1950)	Ukraine (Molloy et al. unpublished data; Yuryshynets et al. 2003)
		Poland (Dobrzanska 1958; Raabe 1956)
		Denmark (Fenchel 1965)
		Russia (Laruelle et al. 1999)
		Belarus (Laruelle et al. 1999)
		France (Molloy et al. unpublished data)
		Germany (Molloy et al. unpublished data)
		Greece (Molloy et al. unpublished data)
		Italy (Molloy et al. unpublished data)
		The Netherlands (Molloy et al. unpublished data)
		Ukraine (Molloy et al. unpublished data)
Sphenophrya dreissenae	P	Poland (Dobrzanska 1958)	Ukraine (Yuryshynets et al. 2003)
		Russia (Laruelle et al. 1999)
		France (Minguez et al. 2009)
		Belarus (Molloy et al. unpublished data)
		Denmark (Molloy et al. unpublished data)
		Germany (Molloy et al. unpublished data)
		Italy (Molloy et al. unpublished data)
		The Netherlands (Molloy et al. unpublished data)
		Ukraine (Molloy et al. unpublished data)
Ophryoglena hemophaga	P	Russia (Fokin et al. 2003; Molloy et al. 1996)	Russia (Pryanichnikova et al. 2011; Tyutin and Scherbina 2006)
		Germany (Fokin et al. 2003)
		The Netherlands (Fokin et al. 2003)
		France (Minguez and Giambérini 2012; Minguez et al. 2009, 2013)
		Ireland (Burlakova et al. 2006a)
		Poland (Yuryshynets 2009)
		Lithuania (Chuševė et al. 2012)
		Denmark (Molloy et al. unpublished data)
		Switzerland (Molloy et al. unpublished data)
		Ukraine (Molloy et al. unpublished data)
Ophryoglena sp. (small form)	P	Belarus (Karatayev et al. 2000a, 2002b, 2003a) France (Minguez et al. 2013)
Ancistrumina limnica	C	Poland (Raabe 1956)	Ukraine (Molloy et al. unpublished data; Yuryshynets 2019)
		The Netherlands (Laruelle et al. 1999)
		Greece (Laruelle et al. 1999)
		Belarus (Karatayev et al. 2000a)
		Ireland (Burlakova et al. 2006a)
		Russia (Molloy et al. unpublished data)
		Ukraine (Molloy et al. unpublished data)
Peritrichia	C	Belarus (Molloy et al. unpublished data)
		Greece (Molloy et al. unpublished data)
		The Netherlands (Molloy et al. unpublished data)
		Ukraine (Molloy et al. unpublished data)
Trematodes
Bucephalus polymorphus	P	France (de Kinkelin et al. 1968b; Minguez and Giambérini 2012; Wallet and Lambert 1986)	Ukraine (Chernogorenko and Boshko 1992)
		Poland (Baturo 1977; Kulczycka 1939; Taskinen et al. 2021)
		Russia (Golikova 1960; Kuperman et al. 1994; Pryanichnikova et al. 2011; Tyutin et al. 2005, 2013a)
		Ukraine (Chernogorenko and Boshko 1992; Zdun 1965)
		Kazakhstan (Smirnova and Ibrasheva 1967)
		Uzbekistan (Aristanov 1986, 1992)
		Belarus (Karatayev et al. 2000a; Stunžėnas et al. 2004)
		Croatia (Lajtner 2012; Lajtner et al. 2008)
		Czech Republic (Ondračková et al. 2015)
		Denmark (Molloy et al. unpublished data)
		Germany (Molloy et al. unpublished data)
		The Netherlands (Molloy et al. unpublished data)
Phyllodistomum macrocotyle	P	The Netherlands (Kraak and Davids 1991)
		Poland (Kulczycka 1939; Petkevičiūtė et al. 2015; Travina et al. 2021; Wisniewski 1957)
		Belarus (Karatayev 1983; Karatayev et al. 2000a; Lyakhnovich et al. 1983; Petkevičiūtė et al. 2015; Stunžėnas et al. 2004; Travina et al. 2021)
		Russia (Kuperman et al. 1994; Molloy et al. 1996; Petkevičiūtė et al. 2015; Travina et al. 2019, 2021)
		Ukraine (Zdun 1965)
		Kazakhstan (Smirnova and Ibrasheva 1967)
		Spain (Peribáñez et al. 2006, 2011)
		France (Minguez and Giambérini 2012; Minguez et al. 2013)
		Lithuania (Petkevičiūtė et al. 2015, 2020; Travina et al. 2021)
		Germany (Molloy et al. unpublished data)
		Greece (Molloy et al. unpublished data)
		Italy (Molloy et al. unpublished data)
Echinoparyphium recurvatum	P	Russia (Ginezinskaja 1959; Kochnev 1977; Pryanichnikova et al. 2011)
		Belarus (Mastitsky and Veres 2010)
		Croatia (Lajtner 2012)
Echinoparyphium echinatoides	P	Ukraine (Chernogorenko and Boshko 1992)
		Russia (Pryanichnikova et al. 2011; Tyutin et al. (2013a)
Echinostoma paraulum	P	Russia (Kochnev 1977)
Echinostomatidae (unidentified)	P	Ukraine (Yuryshynets 1999, 2019)	Russia (Pryanichnikova et al. 2011)
		France (Minguez and Giambérini 2012)	Ukraine (Yuryshynets 1999, 2019)
		Belarus (Karatayev et al. 2000a)
		Denmark (Molloy et al. unpublished data)
		Germany (Molloy et al. unpublished data)
		Greece (Molloy et al. unpublished data)
		Italy (Molloy et al. unpublished data)
		The Netherlands (Molloy et al. unpublished data)
Leucochloridiomorpha constantiae	P	Ukraine (Chernogorenko and Boshko 1992; Molloy et al. 1997)
Sanguinicola sp.	P	Poland (Stanczykowska 1977)
Aspidogaster limacoides	P	Russia (Kuperman et al. 1994; Molloy et al. 1996; Nagibina and Timofeeva 1971; Pryanichnikova et al. 2011; Tyutin and Scherbina 2006; Tyutin et al. 2013a; Zhokhov 2001)	Russia (Popova and Biochino 2001)
		Croatia (Lajtner 2012)
Aspidogaster conchicola	P	Poland (Kulczycka 1939)
		Ukraine (Chernogorenko and Boshko 1992)
Aspidogaster (unidentified)	P	Belarus (Karatayev et al. 2000a)
		France (Minguez et al. 2011)
		The Netherlands (Molloy et al. unpublished data)
Haplosporidians
Haplosporidium raabei	P	The Netherlands (Bowmer and van der Meer 1991; de Kock and Bowmer 1993; Molloy et al. 2012)
		Germany (Molloy et al. unpublished data)
		France (Molloy et al. unpublished data)
Prokaryote bacteria
Rickettsiales-like and/or Chlamydiales-like organisms	P	Greece (Molloy et al. 2001) France (Minguez and Giambérini 2012; Minguez et al. 2012, 2013) United States (Minguez et al. 2013)
Nematodes
Brevitobhlus stefanskii	C	Belarus (Mastitsky and Gagarin 2004)
Chromadorita leuckarti	C	Belarus (Mastitsky and Gagarin 2004)
Chromadorina bioculata	C	Belarus (Karatayev et al. 2003a; Mastitsky and Gagarin 2004)
	C	Sweden (Mastitsky et al. 2008)
Crocodorylaimus flavomaculatus	C	Belarus (Mastitsky and Gagarin 2004)
Dorylaimus stagnalis	C	Belarus (Mastitsky and Gagarin 2004)
	C	Russia (Kuperman et al. 1994)
Epitobrilus medius	C	Belarus (Mastitsky and Gagarin 2004)
Eumonhystera pseudobulbosa	C	Belarus (Mastitsky and Gagarin 2004)
Eumonhystera vulgaris	C	Belarus (Mastitsky and Gagarin 2004)
Laimydorus sp.	C	Sweden (Mastitsky et al. 2008)
Monhystera uncispiculatum	C	Belarus (Mastitsky and Gagarin 2004)
Monhystera paludicola	C	Belarus (Mastitsky and Gagarin 2004)
Monhystera iemani	C	Belarus (Mastitsky and Gagarin 2004)
Monhystrella sp.	C	Belarus (Karatayev et al. 2003a)
Mononchus truncatus	C	Belarus (Mastitsky and Gagarin 2004)
Neoactinolaimus dzjubani	C	Belarus (Mastitsky and Gagarin 2004)
Plectus cirratus	C	Belarus (Mastitsky and Gagarin 2004)
Plectus palustris	C	Belarus (Mastitsky and Gagarin 2004)
Punctodora ratzeburgensis	C	Belarus (Mastitsky and Gagarin 2004)
Rhabdolaimus terrestris	C	Belarus (Mastitsky and Gagarin 2004)
Semitobrilus gagarini	C	Belarus (Mastitsky and Gagarin 2004)
Tobrilus gracilis	C	Belarus (Mastitsky and Gagarin 2004)
Tobrilus helveticus	C	Belarus (Mastitsky and Gagarin 2004)
Tobrilus tenuicaudatus	C	Belarus (Karatayev et al. 2003a)
Tridentulus floreanae	C	Belarus (Karatayev et al. 2003a; Mastitsky and Gagarin 2004)
Tripyla glomerans	C	Belarus (Mastitsky and Gagarin 2004)
Nematoda (unidentified)	C	Russia (Kuperman et al. 1994; Pryanichnikova et al. 2011)	Russia (Tyutin et al. 2013a)
		Belarus (Karatayev et al. 2000a, 2003a)
		Ireland (Burlakova et al. 2006a)
		Sweden (Mastitsky et al. 2008)
		Lithuania (Chuševė et al. 2012)
		Poland (Krasutska 2017)
		Czech Republic (Molloy et al. unpublished data)
		Denmark (Molloy et al. unpublished data)
		France (Molloy et al. unpublished data)
		Germany (Molloy et al. unpublished data)
		Greece (Molloy et al. unpublished data)
		Italy (Molloy et al. unpublished data)
		The Netherlands (Molloy et al. unpublished data)
		Switzerland (Molloy et al. unpublished data)
		Ukraine (Molloy et al. unpublished data)
Oligochaetes
Chaetogaster limnaei	P?	Ukraine (Chernogorenko and Boshko 1992)
		Belarus (Karatayev et al. 2000a)
		Germany (Molloy et al. unpublished data)
Psammoryctides baebatus	C	Russia (Kuperman et al. 1994)
P. moldavensis	C	Russia (Kuperman et al. 1994)
		Russia (Kuperman et al. 1994)
Oligochaeta (unidentified)	C	Belarus (Karatayev et al. 2000a)	Russia (Pryanichnikova et al. 2011; Tyutin et al. 2013a)
		Ireland (Burlakova et al. 2006a)	Ukraine (Yuryshynets 2019)
		Denmark (Molloy et al. unpublished data)
		Ukraine (Molloy et al. unpublished data)
Leeches
Caspiobdella fadejewi	C	Russia (Kuperman et al. 1994)
Helobdella stagnalis	C	Russia (Kuperman et al. 1994)
Helobdella sp.	C	Belarus (Karatayev et al. 2000a)	Tyutin et al. 2013a
Erpobdella octoculata	C	Belarus (Karatayev et al. 2000a)
Erpobdella sp.	C		Tyutin et al. 2013a
Chironomids
Chironomus bathophilus	C	Russia (Kuperman et al. 1994)	Ukraine (Yuryshynets 2019)
Cryptochironomus sp.	C	Belarus (Mastitsky and Samoilenko 2005)
Glyptotendipes gr. gripekoveni	C	Belarus (Mastitsky and Samoilenko 2005)
Limnochironomus gr. nervosus	C	Belarus (Mastitsky and Samoilenko 2005)
Limnochironomus tritomus	C	Belarus (Mastitsky and Samoilenko 2005)
Paratanytarsus gr. lauterborni	C	Belarus (Mastitsky and Samoilenko 2005)
Polypedilum gr. convictum	C	Belarus (Mastitsky and Samoilenko 2005)
Tanytarsus gr. gregarius	C	Belarus (Mastitsky and Samoilenko 2005)
Corynoneura celeripes	C	Belarus (Mastitsky and Samoilenko 2005)
Cricotopus gr. algarum	C	Belarus (Mastitsky and Samoilenko 2005)
Cricotopus gr. silvestris	C	Belarus (Mastitsky and Samoilenko 2005)
Cricotopus sp.	C	Belarus (Mastitsky and Samoilenko 2005)
Eukieferiella bicolor	C	Belarus (Mastitsky and Samoilenko 2005)
Prodiamesa sp.	C	Belarus (Mastitsky and Samoilenko 2005)
Orthocladiinae spp.	C	Belarus (Mastitsky and Samoilenko 2005)
Chironomidae (unidentified)	C	Belarus (Karatayev et al. 2000a)
		Ireland (Burlakova et al. 2006a)
		Sweden (Mastitsky et al. 2008)
		Russia (Pryanichnikova et al. 2011)
		Czech Republic (Molloy et al. unpublished data)
		Denmark (Molloy et al. unpublished data)
		Germany (Molloy et al. unpublished data)
		Greece (Molloy et al. unpublished data)
		Ukraine (Molloy et al. unpublished data)
Mites
Unionicola sp.	P?	Ukraine (Chernogorenko and Boshko 1992)
		Russia (Kuperman et al. 1994)	Russia (Tyutin et al. 2013a)
Hydrachnidia (unidentified)	P?	Belarus (Karatayev et al. 2000a)
		Sweden (Mastitsky et al. 2008)

This list does not include C. klimentinus and S. naumiana found exclusively in D. carinata in Lake Ohrid.

Table 11. Endosymbionts reported from the attached Dreissena polymorpha and D. r. bugensis populations in North America.

Endosymbionts	Type of symbiont	D. polymorpha	D. r. bugensis
Ciliates
Ophryoglena hemophaga	P	New York (Molloy unpublished data; Mastitsky personal communication)
Trematodes
Echinostomatidae (unidentified)	P	New York (Karatayev et al. 2012)	New York (Karatayev et al. 2012)
Sphaeridiotrema sp.	P	New York (Karatayev et al. 2012)	New York (Karatayev et al. 2012)
Aspidogaster conchicola	P	New York (Toews et al. 1993)
Nematodes
Achromadora sp.	C		California (Reid et al. 2012)
Chromadorina bioculata	C		California (Reid et al. 2012)
Dichromadora sp.	C		California (Reid et al. 2012)
Diplogaster sp.	C		California (Reid et al. 2012)
Ironus sp	C		California (Reid et al. 2012)
Mononchus sp.	C	New York (Conn et al. 1994)
Monhystrella sp.	C		California (Reid et al. 2012)
Laimydorus sp.	C		California (Reid et al. 2012)
Plectus geophilus	C		California (Reid et al. 2012)
Rhabdolaimus sp.	C		California (Reid et al. 2012)
Seinura sp.	C		California (Reid et al. 2012)
Nematoda (unidentified)	C	New York (Conn et al. 1994; Toews et al. 1993)	New York (Conn et al. 1994)
Oligochaets
Chaetogaster limnaei	P?	New York (Conn et al. 1994, 1996)	New York (Conn et al. 1994, 1996)
Ophidonais serpentine	C	New York (Conn et al. 1994)
Oligochaeta (unidentified)	C	New York (Conn et al. 1994)	New York (Conn et al. 1994)
Chironomids
Paratanytarsus sp.	C	New York (Conn et al. 1994; Ricciardi 1994)	New York (Conn et al. 1994; Ricciardi 1994)
Mites
Hydrachnidia (unidentified)	P?	New York (Conn et al. 1994)

Figure 2. Ciliates from Dreissena spp.: A, Conchophthirus acuminatus; B, Conchophthirus klimentinus (both adapted from Raabe 1971); C, Sphenophrya naumiana; D, Sphenophrya dreissenae (both adapted from Raabe 1966); E, Hypocomagalma dreissenae (adapted from Fenchel 1965); F, Ancistrumina limnica (adapted from Raabe 1947); G, H, trophont and theront of Ophryoglena hemophaga (adapted from Molloy et al. 2005).

Figure 3. Hypothetical life cycle of Ophryoglena hemophaga (adapted from: Canella and Rocchi-Canella 1976; Morton 1993; Owen 1955; Thorp and Covich 1991). In laboratory studies (Molloy et al. 2005), (1) the presence of “trophonts” (the feeding stage) was repeatedly observed in the ducts of the digestive gland (right image); (2) well-fed trophonts were subsequently commonly found to have recently emerged as “protomonts” from infected D. polymorpha (left image), but it is unknown if they emerged via the exhalant siphon (as pictured) or the inhalant siphon; (3) likewise, protomont encystment into the “tomont” stage, followed by “tomite” production within the tomont, followed by the exit of these tomites from the cyst as free swimming “theronts” was also commonly observed, but it is unknown if these theronts immediately initiate reinfection by entering through the inhalant siphon (as pictured) or possibly go through another encystment-tomont-tomite-theront production cycle again outside the mussel before initiating infection again.

Figure 4. Life cycle of Bucephalus polymorphus (adapted from Molloy et al. 1997).

Figure 5. Life cycle of Phyllodistomum macrocotyle (adapted from Molloy et al. 1997 with information in Petkevičiūtė et al. 2020).

Figure 6. Sporocysts (yellow bodies) within the gills of a Phyllodistomum sp. in Dreissena carinata from Lake Ohrid, Republic of North Macedonia (Credit: D. P. Molloy).

Figure 7. Life cycle of Echinoparyphium recurvatum (adapted from Molloy et al. 1997).

Figure 8. Histological section of advanced infection with the haplosporidian Haplosporidium raabei in gill of Dreissena polymorpha (adapted from Molloy et al. 2012).

Table 12. Competitors of Dreissena spp. reported from Europe (E) and North America (NA).

Competitor	Conti-nent	Dreissena species, location, references
Sponges
Ochridaspongia rotunda	E	D. carinata, Macedonia (Arndt 1937)
Spongilla carteri	E	D. polymorpha, Hungary (Sebestyén 1937)
S. lacustris	NA	D. polymorpha, Quebec, Ohio (Ricciardi et al. 1995)
	NA	D. r. bugensis, Quebec, Ohio (Ricciardi et al. 1995)
Eunapius (=Spongilla) fragilis	E	D. polymorpha, Hungary (Sebestyén 1937)
	NA	D. polymorpha, Quebec, Ohio (Ricciardi et al. 1995)
	NA	D. polymorpha, Indiana (Early and Glonek 1999; Lauer 1997; Lauer and Spacie 2000, 2004)
	NA	D. r. bugensis, Quebec, Ohio (Ricciardi et al. 1995)
Ephydatia fluviatilis	E	D. polymorpha, Poland (Piesik 1983)
	E	D. polymorpha, Italy (Lancioni and Gaino 2009)
Ephydatia muelleri	NA	D. polymorpha, Quebec, Ohio (Ricciardi et al. 1995)
	NA	D. polymorpha, Indiana (Lauer 1997; Lauer and Spacie 2000)
	NA	D. r. bugensis, Quebec, Ohio (Ricciardi et al. 1995)
Bryozoans
Plumatella repens	E	D. polymorpha, Poland (Piesik 1983)
Pectinatella magnifica	NA	D. polymorpha, New York (Conn and Conn 1993)
Lophopodella carteri	NA	D. polymorpha, Indiana (Lauer et al. 1999)
Other competitors
Chlorophytes
Cladophora sp.	E	D. polymorpha, Hungary (Sebestyén 1937)
	E	D. polymorpha, The Netherlands (Smit et al. 1993)
Crustaceans
Chelicorophium curvispinum	E	D. polymorpha, Hungary (Sebestyén 1937)
	E	D. polymorpha, The Netherlands (Paffen et al. 1994; Smit et al. 1993; van den Brink et al. 1993; van der Velde et al. 1994)
		D. polymorpha, Germany (Jantz and Schöll 1998)
Hydrozoans
Cordylophora caspia	E	D. polymorpha, The Netherlands (Smit et al. 1993)
	E	D. polymorpha, Poland (Piesik 1983)
Bivalves
Mytilaster lineatus	E	D. elata, Caspian Sea (Karpinsky 2010; Karpinsky et al. 2005; Logvinenko and Starobogatov 1968)
	E	D. caspia, Caspian Sea (Karpinsky 2010; Karpinsky et al. 2005; Logvinenko and Starobogatov 1968)
D. r. bugensis	E, NA	D. polymorpha, multiple waterbodies in Europe and North America (Balogh et al. 2018; bij de Vaate et al. 2014; Heiler et al. 2013; Hetherington et al. 2019; Karatayev and Burlakova 2022a, 2022b; Karatayev et al. 2011a, 2015, 2021a, 2021b; Noordhuis et al. 2016; Orlova et al. 2004, 2005; Strayer et al. 2019)

Table 13. Total number of Dreissena spp. natural enemies recorded in Europe and North America according to Molloy et al. (1997) and current study (2022).

Natural enemy	Europe		North America		Both continents		Total
	1997	2022	1997	2022	1997	2022	1997	2022
Fish	30	42	19	39	3	4	46	77
Birds	21	22	20	22	5	5	36	39
Crayfish	3	10	3	4	0	0	6	14
Others	5	7	3	7	1	1	7	13
All Predators	59	81	45	72	9	10	95	143
Parasites	17	20	3	6	3	5	17	21
Commensals	15	55	2	15	0	5	17	65
All endosymbionts	32	75	5	21	3	10	34	86
Competitors	8	10	5	6	2	2	11	14
All natural enemies	99	166	55	99	14	22	140	243

This list includes C. klimentinus and S. naumiana found exclusively in D. carinata in Lake Ohrid. The total number of predators and competitors in Molloy et al. (1997) were reported erroneously as 176 and 10.

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