Sander lucioperca (Pikeperch): Dossier

Distribution

IUCN Red List status

least concern

Information

Authors: Jenny Volstorf, Tanya Slosberg
Version: B | 1.1 (2022-01-22)

Reviewer: Caroline Marques Maia
Editor: Billo Heinzpeter Studer

Initial release: 2016-05-15
Version information:

Appearance: B
Last minor update: 2022-01-22

Cite as: »Volstorf, Jenny, and Tanya Slosberg. 2022. Sander lucioperca (Dossier). In: fair-fish database, ed. fair-fish. World Wide Web electronic publication. First published 2016-05-15. Version B | 1.1. https://fair-fish-database.net.«

1 Remarks

1.1 General remarks

Escapees and consequences: negative or at most unpredictable for the local ecosystem

Unpredictable influence:
- WILD: proportion of Sander lucioperca in riverine sites in UK increased from 0.1% in 1961 to 0.5% in 2000 1.
Competition:
- WILD: 23 years after introduction into lake Egirdir, Turkey, S. lucioperca was only species with large stock. Cyprinus carpio and Vimba vimba populations had decreased sharply. In 2006, Phoxinellus anatolicus, Orthrias angorae, Semi-noemacheilus lendli, Hemmigramocapoeta kemali were found in small numbers. Phoxinellus egirdiri, Phoxinellus handlirschi, Phoxinellus zeregi probably extinct 2.
Disease transmission: no data found yet.
Interbreeding: no data found yet.

1.2 Other remarks

No data found yet.

2 Ethograms

In the wild: on feeding, daily rhythm, migration, cooperation

For feeding ➝ 3.
For daily rhythm ➝ 4 5 6.
For migration ➝ 7 8 9.
For cooperation ➝ 4.

In the farm or lab: on feeding, reproduction

For feeding ➝ 10.
For courtship and spawning ➝ 11.

3 Distribution

Natural distribution: Europe, inland and coastal waters

Observations Scandinavia: lake Hiidenvesi, Finland 12, lake Hjälmaren, Sweden 7, lake Oulujärvi, Finland 13, river Oulujoki, Finland 9.
Observations continental Europe: Aras lake dam, Azerbaijan 14, Curonian lagoon, Baltic Sea, Lithuania 8, Elbe river, Czech Republic 15, Fumemorte canal, Rhone delta, southern France 16, Lipno reservoir, Czech Republic 17, Talty lake, Mazurian lake land, Poland 18, Volga river, Russia 19.

Introduced: Europe, inland waters

Observations England: Relief and Cut-off Channel, England 4, Stanborough lake, Hertfordshire, and Ash Drain (Great Ouse Middle Level), Cambridgeshire, England 20.
Observations France: Castillon and Treignac reservoirs, France 21.
Observations Turkey: lake Egirdir, Turkey 22, various inland waters, Turkey 2, Seyhan Dam lake, Turkey 23.

4 Natural co-existence

No data found yet.

5 Substrate and/or shelter

5.1 Substrate

Substrate range, substrate preference: lives over sand and fine substrate

Plants:
- WILD: in summer, moved in open water without vegetation: river Oulujoki, Finland 9.
- LAB: in 80 x 25 x 0.8 m pond with sandy bottom, single ADULTS in 3 x 4 m enclosures of clear PVC and wooden frame. Higher predation rate without vegetation (ca 1.25-1.75 versus 0-0.25 prey fish/d) than with artificial vegetation of green buoyant polypropylene strings attached to PVC plate 3.
Rocks and stones: no data found yet.
Sand and mud:
- WILD: JUVENILES were found over fine substrate without mud: Lipno reservoir, Czech Republic 17.
- WILD: was found over sands, slabs, cobbles: Castillon reservoir, France (introduced) 21.
- For substrate and nest building ➝ D1.
Other substrate: no data found yet.

5.2 Shelter or cover

Shelter or cover preference: uses shelter during hunting (further research needed)

Plants: no data found yet.
Rocks and stones: no data found yet.
Sand and mud: no data found yet.
Other cover: no data found yet.
For shelter and hunting ➝ D2.

6 Food, foraging, hunting, feeding

6.1 Trophic level and general considerations on food needs

Trophic level: 4.0

Observations: 4.0±0.8 se 24.

Impacts of feed fishery: contributes to overfishing, challenges animal welfare

Carnivorous D3. The fishery that provides fish meal and fish oil has two major impacts:
1. It contributes considerably to overfishing, as it accounts for 1/4 25 or even 1/3 26 of the world catch volume.
2. It challenges animal welfare, because in the face of 450-1,000 MILLIARD wild fishes caught worldwide each year to be processed into fish meal or fish oil 27, the individual fish gets overlooked and, thus, suffering increases at rearing, live marketing, and slaughtering levels 28.

6.2 Food items

Food items, food preference: carnivorous, cannibalistic at times; increasing prey size with increasing age

Food items: carnivorous, cannibalistic at times:
- Observations WILD: mainly fish, followed by invertebrates (mainly Amphipoda): Relief and Cut-off Channel, England (introduced) 31-4, Curonian lagoon, Baltic Sea, Lithuania and Baltic Sea, Lithuania 8, mainly fish, e.g. smelt Atherina boyeri, Mugil spp., pumpkinseed Lepomis gibbosus, prussian carp Carassius gibelio, bleak Alburnus alburnus, Abramis spp., stone moroko Pseudorasbora parva, then crustaceans, no cannibalism: Fumemorte canal, Rhone delta, southern France 16, lake Egirdir, Turkey (introduced) 22, mainly smelt Osmerus eperlanus and other fish (roach Rutilus rutilus, bream Abramis brama, bleak Alburnus alburnus, perch Perca fluviatilis, ruffe Gymnocephalus cernuus), 4.8% JUVENILES of ≥1 year cannibalistic in two of 12 months observation, July and September, at highest abundance of young JUVENILES: lake Hiidenvesi, Finland 12, JUVENILES: mainly cyclopoid copepods during the day, cladocerans Daphnia sp. during the night: Lipno reservoir, Czech Republic 17.
Food items and habitat: no data found yet.
Food items and life stages: JUVENILES mainly crustaceans, insects, small fishes, ADULTS mainly larger fishes, cannibalistic at times:
- WILD: Curonian lagoon, Baltic Sea, Lithuania 8:
  JUVENILES (>1 year): mainly mysids Mysidae,
  ADULTS (3-4 years): mainly ruffe Gymnocephalus cernuus, roach Rutilus rutilus, perch Perca fluviatilis.
- WILD: Baltic Sea, Lithuania 8:
  JUVENILES (>1 year): mainly smelt Osmerus eperlanus, mysids Mysidae, goby Pomatoschistus microps,
  ADULTS (3-4 years): mainly Atlantic herring Clupea harengus, sand eel Ammodytes tobianus, smelt Osmerus eperlanus.
- WILD, JUVENILES-ADULTS: 15-34 cm: mainly fish (mainly Knipowitschia sp. and Aphanius anatoliae anatoliae, 0.6% cannibalism): lake Egirdir, Turkey (introduced) 22:
  15-24 cm: fish, odonats Calopteryx splendens, mysids Mysis, amphipods Gammarus,
  25-29 cm: fish, odonats Calopteryx splendens, frogs Rana, mysids Mysis, dipterans Chironomus,
  30-34 cm: fish, frogs Rana, odonats Calopteryx splendens.
- WILD: Castillon reservoir, France (introduced) 21:
  <25 cm: mainly insects (Diptera, Plecoptera, Trichoptera, Ephemeroptera), fish (bleak Alburnus alburnus, pikeperch S. lucioperca, perch Perca fluviatilis), crustaceans (Isopodia)
  25-33 cm: mainly fish (bleak Alburnus alburnus, pikeperch S. lucioperca, perch Perca fluviatilis, roach Rutilus rutilus), insects (Diptera), crustaceans (Isopodia),
  >33 cm: mainly fish (bleak Alburnus alburnus, pikeperch S. lucioperca, bream Abramis brama, perch Perca fluviatilis, roach Rutilus rutilus, brown trout Salmo trutta).
- WILD: Treignac reservoir, France (introduced) 21:
  <25-33 cm: mainly insects (Trichoptera, Diptera, Plecoptera) and fish (perch Perca fluviatilis, roach Rutilus rutilus, pikeperch S. lucioperca),
  >33 cm: mainly fish (pikeperch S. lucioperca, roach Rutilus rutilus), insects (Diptera, Trichoptera).
Food preference: no data found yet.
Food partitioning: no data found yet.
Prey density:
- LAB: single JUVENILES (192-228 mm) in pool of 2.4 m diameter, depth 35 cm with 1, 2, 4, 8, 16 prey (within subject). With increasing prey density, no tendency in number of attacks taken until capture and no tendency in time to first foraging move (median 72 s, mean 271 s), but tendency of decreasing time to first attack. Capture success in 18 of 211 attacks, 8.5%. No difference in capture success when attacking single or multiple prey. Results indicate no confusion by increasing prey density 10.
Prey size selectivity:
- LAB: in 80 x 25 x 0.8 m pond with sandy bottom, single ADULTS in 3 x 4 m enclosures of clear PVC and wooden frame either presented with crucian carp Carassius carassius (of 80% 85-115 mm), rudd Scardinius erythrophthalmus (of 80% 85-140 mm), or combination of both species. No difference in predation rate of crucian carp when presented alone or together with rudd (ca 1.25-1.75 fish/d). Lower predation rate of rudd when presented together with carp than alone (0.25 prey fish/d versus 1.25 prey fish/d). Overall, no selection of smaller size, though 3.
Particle size: no data found yet.

6.3 Feeding behaviour

Feeding style, foraging mode: predator, hunter

WILD: wandering predator: actively pursued prey instead of ambushing 3.
LAB: single JUVENILES (192-228 mm) in pool of 2.4 m diameter, depth 35 cm with two shelters of parallel bricks (25 x 12 cm) in pool centre. JUVENILES actively chased and attacked single and schools of prey, one sit-and-wait attack. Usually one attack per chase. Predation cycle consisting of start to first move, first move to attack, attack to capture lasted ca 21 min. In 16 of 35 trials, JUVENILES hid in shelters before attacks 10.
For hunting and vision ➝ D4.

Effects on feeding: direct relation with temperature

Feeding and temperature:
- WILD, JUVENILES-ADULTS: with decreasing water temperatures, increasing rate of empty stomachs: lake Egirdir, Turkey (introduced) 22.
- FARM, FRY: poor food uptake and growth at <18-20 °C 32.

7 Photoperiod

7.1 Daily rhythm

Daily rhythm: individual differences: crepuscular, nocturnal, diurnal

Daily rhythm:
- WILD: 0+ JUVENILES were found with stomach content during day and night: Lipno reservoir, Czech Republic 17.
- WILD, JUVENILES-ADULTS: predation from dusk till dawn: Relief and Cut-off Channel, England (introduced) 4.
- WILD, ADULTS: individual differences: crepuscular, diurnal, and also switching between rhythms over the year: Fumemorte canal, Rhone delta, France 5.
- WILD: ADULTS moved average 125-130 m during dusk and dawn (ca 5-30 lux), average 75 m during day (>30 lux) and night (<5 lux): Elbe river, Czech Republic 6.
- LAB, JUVENILES (192-228 mm): with decreasing light intensity from 500 lux to <1 lux, foraging activity increased. Prey captures in 18 of 35 trials during dim light, another 13 during night 10 (for details on the study ➝ D3 "Prey density").
Nocturnal activity: ➝ Daily rhythm.
Phototaxis: no data found yet.
For PHOTOPERIOD and spawning ➝ D5.

Photoperiod and stress: 24 h darkness is stressful (further research needed)

LAB: two mixed-sex pairs of ADULTS in 50 cm deep 1,500 L tanks at density 0.9 kg/m³ under lighting regimes of either "12 h light and 12 h dark", 24 h light, or 24 h dark for 40 days. Half of the pairs were stressed by confining ADULTS to 10 cm water level, capturing them, and holding them outside the water for 20 s each day. No difference in cortisol levels indicating adaptation to long-term stress, but higher glucose level in unstressed females in 24 h dark compared to stressed females (ca 190 mg/dL versus 90 mg/dL). No difference in hematocrit and hemoglobin, but higher amount of white blood cells in unstressed females in 24 h dark compared to stressed females (ca 22,000 cells/mm versus 13,000 cells/mm) 33.

7.2 Light intensity

Light intensity preference: 1 lux (further research needed)

Light intensity preference:
- LAB: individual 0+ and 1+ JUVENILES in 1 m³ tanks divided into four compartments covered with different amounts of white paper. On day 1 with 304 lux in all compartments, no difference in preference. On day 2, no difference in preference between compartments with 304, 149, 92, and 26 lux. On days 3 and 4, 0+ and 1+ JUVENILES preferred the compartment with 26 lux over the compartments with higher light intensities (75-100% versus 0-20% time spent).
  In second experiment, on day 1 with 55 lux in all compartments, no difference in preference. On day 2, no difference in preference between compartments with 55, 21, 12, and 1 lux. On days 3 and 4, 0+ and 1+ JUVENILES preferred the compartment with 1 lux over the compartments with higher light intensities (75-100% versus 0-25% time spent) 34.

Light intensity and growth: inverse relation (further research needed)

LAB: JUVENILES in 1 m³ rearing tanks at stocking density 0.9 kg/m³ and light intensity of either 385.7 lux or 45.1 lux. After 56 days, higher weight under low (45.9 g versus 40.6 g) than under high intensity, but hardly noticeable difference in specific growth rate (3.6%/d versus 3.4%/d) and FOOD CONVERSION RATIO (1.0 versus 1.1). JUVENILES under high light intensity hid beneath feeder to shelter from light, JUVENILES under low light intensity spread evenly in water. After pause of 10 days, second experiment with stocking density 3.2 kg/m³ and either 8.0 lux or 1.2 lux. After 56 days, higher weight under low (113 g versus 109 g) than under high intensity, but no difference in specific growth rate (1.7-1.8%/d) and FOOD CONVERSION RATIO (0.8) 35.
For light intensity and foraging ➝ D6.

7.3 Light colour

Light colour and growth: highest under red light (further research needed)

LAB: single JUVENILES in 40 x 20 x 25 cm 15 L aquaria covered with blue (435 nm), red (610 nm), yellow (564 nm), green (534 nm), or white filter and lined with paper of same colour at 50 lux. After 42 days, no difference in feed intake (10.7-12.3 g/kg/d), but higher final weight (106.1 g versus 72.1 g) and higher specific growth rate (2.4%/d versus 1.6%/d) under red than white light; other light colours in between. No effect of colour on cortisol levels due to large individual differences. Results indicate probable habituation to eutrophic environment which light of wavelength <600 nm hardly penetrates 36.

8 Water parameters

8.1 Water temperature

Standard temperature range, temperature preference: 0-27 °C, capable of surviving under ice cover (further research needed)

Standard temperature range:
- Observations WILD: 8-24 °C: lake Hjälmaren, Sweden 7, ca 2-20 °C: Curonian lagoon and Baltic Sea, Lithuania 8, summer-autumn 10-20 °C, winter (Nov-April) ice cover: river Oulujoki, Finland 9, 1-27 °C: Fumemorte canal, Rhone delta, France 5, 4-5 °C in winter, 25-26 °C in summer: lake Egirdir, Turkey (introduced) 22, 0-24 °C: Elbe river, Czech Republic 15, 15-17 °C: Lipno reservoir, Czech Republic 17, maximum surface temperature: 22-25 °C: Castillon and Treignac reservoirs, France (introduced) 21.
- FARM: JUVENILES overwintered 176 days in 0.8 m deep pond ice-covered on 100 days. No difference in survival rate (83.3%) between group of 31.6 g weight and group of 13.8 g weight 37.
Temperature preference: no data found yet.
For temperature and...
...feeding ➝ D7,
...(swimming) activity ➝ D8,
...water velocity ➝ D9,
...depth ➝ D10.

Temperature and growth: optimally 23 °C (further research needed)

Temperature must exceed: no data found yet.
Temperature must not go beyond: no data found yet.
Optimal temperature for growth:
- FARM: JUVENILES in 500 L tanks at either 6, 9, 12, 15, 23 °C. After five months, higher weight (ca 750 g versus 400 g) and higher length (ca 42.5 cm versus 35 cm) under 23 °C than under the lower temperatures 38.
For temperature and feeding ➝ D7.

8.2 Oxygen

Dissolved oxygen range: 5.5-12.9 mg/L

Observations WILD: 5.5-12.9 mg/L Elbe river, Czech Republic 15, 8-11 mg/L: Lipno reservoir, Czech Republic 17.

8.3 Salinity

Salinity tolerance, standard salinity range: probably euryhaline (further research needed), 0.03-10+ ppt

Salinity tolerance:
- Natural and introduced distribution in fresh water or brackish water at most D11 D12 D13.
Standard salinity range: 0.03-10+ ppt:
- Observations WILD: 0.03-1.6 ppt Curonian lagoon, Baltic Sea, Lithuania 8, 0.1 ppt in summer, 5 ppt in winter: Fumemorte canal, Rhone delta, France, >10 ppt: Vaccarès lagoon, France 5.

Salinity and growth: depends on water temperature (further research needed)

LAB: wild-caught 0+ JUVENILES in groups of five in 100 L glass aquaria. After four weeks in 17-21 °C water, higher weight increment at 5 ppt (ca 23 g versus 9 g) than 0 ppt, 2 ppt in between. After six weeks in 9-12 °C water, no difference in weight increments at 0, 2, or 5 ppt (ca 16-19 g) 8.

8.4 pH

Standard pH range: 6.5-6.9 (further research needed)

Standard pH range:
- Observations WILD: pH 6.5-6.9: river Oulujoki, Finland 9.
pH preference: no data found yet.

8.5 Turbidity

Standard turbidity range: Secchi depth 0.4-10 m

Standard turbidity range: no data found yet.
Secchi depth (water transparency): 0.4-10 m:
- Observations WILD: 2.5 m: lake Hjälmaren, Sweden 7, 0.4-2.0 m Curonian lagoon, Baltic Sea, Lithuania 39-8, >40 cm: Fumemorte canal, Rhone delta, France 5, <2 m: Treignac reservoir, France, 3-10 m: Castillon reservoir, France (introduced) 21.

8.6 Water hardness

No data found yet.

8.7 NO4

No data found yet.

8.8 Other

No data found yet.

9 Swimming

9.1 Swimming type, swimming mode

Swimming type, swimming mode: sub-carangiform

Observations: 40.

9.2 Swimming speed

Swimming speed: max 1.6 body lengths/s (further research needed)

Absolute swimming speed: no data found yet.
Relative swimming speed:
- Observations WILD, ADULTS: maximum speed during 2 h-interval in March by females: 1.6 body lengths/s: Fumemorte canal, Rhone delta, France 5.
(Swimming) activity and temperature:
- WILD: with decreasing water temperatures, JUVENILES-ADULTS became stationary: river Oulujoki, Finland 9.

Standard velocity range, velocity preference: 0.01-0.9 cm/s, decreasing with decreasing temperatures (further research needed)

Standard velocity range:
- Observations WILD: 0.01-0.9 cm/s, mean 0.2 cm/s: river Oulujoki, Finland 9.
Velocity preference: no data found yet.
Velocity and temperature:
- WILD: with decreasing water temperatures, JUVENILES-ADULTS preferred lower velocities: river Oulujoki, Finland 9.

9.3 Home range

Home range: average 0.2-6.3 km, range 0-7.7 km

WILD, JUVENILES-ADULTS: daily range in summer: almost immobile to 7,652 m, mean 947 m: river Oulujoki, Finland 9.
WILD, ADULTS: females with higher daily range of movement than males: average 905-2,010 m/d versus 168-598 m/d, range in females: 22-3,410 m/d. Daily distance travelled across genders: average 313-3,932 m/d, range 53-7,561 m/d, 54% >1,000 m/d: Fumemorte canal, Rhone delta, France 5.
WILD, ADULTS: more diurnal movement in winter (ca 240 m versus 50-80 m) compared to other seasons, probably to increase predation success given decreased abundance of prey. More longitudinal movement in spring and autumn (ca 2,900 m versus 400 m) compared to winter; stationary in summer: Elbe river, Czech Republic 15.

9.4 Depth

Depth range, depth preference: range 0-30.8 m, moves deeper during the day probably to avoid predators

Depth range in the wild:
- Observations WILD, JUVENILES: caught in depth 1.5-4 m: lake Hiidenvesi, Finland 12, 0-6 m: Lipno reservoir, Czech Republic 17.
- Observations WILD, JUVENILES-ADULTS: 1 m: Fumemorte canal, Rhone delta, France 5.
- Observations WILD, ADULTS: range 1.2-30.8 m, mean 9.8 m: river Oulujoki, Finland 9, 0-2 m: Elbe river, Czech Republic 6.
- For depth and nest building ➝ D1.
Depth in cages or tanks: no data found yet.
Depth preference: no data found yet.
Depth and daily rhythm: moves deeper during the day probably to avoid predators:
- WILD, JUVENILES: 20-40 IND/1,000 m³ of age 0+ in 0-5 m during day, 200-600 IND/1,000 m³ of age 0+ in 0-6 m during night, migrating upwards during dusk and downwards during dawn: Lipno reservoir, Czech Republic 17.
- WILD: ADULTS occurred at 0-1.5 m during dusk or dawn, 0-2 m during day, 0-0.5 m during night: Elbe river, Czech Republic 6.
Depth and low temperatures:
- WILD, JUVENILES-ADULTS: with decreasing water temperatures, especially females moved to deeper water from 7-15 m in summer to 21-23 m in winter: river Oulujoki, Finland 9.
Depth and high temperatures: no data found yet.
Position in habitat and age: no data found yet.
Depth and light intensity: no data found yet.
Depth and noise: no data found yet.
Depth and threat: no data found yet.

9.5 Migration

Migration type: semi-anadromous

JUVENILES are mostly resident, some migrate >9 km during one year:
- Observations WILD: during one year, 10 of 17 JUVENILES-ADULTS remained within 9.2 km of study site; four moved downstream outside study area. 15 of 20 JUVENILES-ADULTS descended downstream summer-autumn (June-October), five overwintered: river Oulujoki, Finland 9.
- WILD: during summer, most JUVENILES were recaptured at average distance of 2.9 km, 50% at distance of 1 km, range 0-32 km. JUVENILES <30 cm were recaptured within 2 km, JUVENILES of 35-39 cm within 4.5 km. During winter, most JUVENILES were recaptured at average 6.3 km, 50% at 6 km, range 0-28 km: lake Hjälmaren, Sweden 7.
ADULTS migrate to the sea after spawning:
- Observations WILD: ADULTS migrated from Curonian lagoon, Baltic Sea, Lithuania to the open Baltic Sea, offshore Lithuania in April, returned in September 8.

10 Growth

10.1 Ontogenetic development

Mature egg: 0.7-0.9 mm

Observations time from fertilisation until hatching: no data found yet.
Observations size WILD: 0.7-0.9 mm diameter: Stanborough lake, Hertfordshire, and Ash Drain (Great Ouse Middle Level), Cambridgeshire, England (introduced) 20.
Observations weight: no data found yet.

Larvae: hatching to 4 days (further research needed)

Observations age at yolk sac absorption LAB: ca at day 4 post hatch 18.
Observations TOTAL LENGTH: no data found yet.
Observations weight: no data found yet.

Fry: beginning of exogenous feeding, 4-48 days, 5.6-15 mm, 0.5-60 mg

Observations beginning of exogenous feeding: no data found yet.
Observations age, TOTAL LENGTH, and weight LAB: 4 days, 5.6 mm, 0.5 mg 18, inflation of swim bladder ca at day 8-18 18, 48 days, 15 mm, 0.06 g 42.

Juveniles, sexual maturity: fully developed (48 days) to beginning of maturity (1-3 years), 4-32.2+ cm, 0.5-553 g

Observations age: ➝ D14.
Observations age, TOTAL LENGTH, and weight FARM: 2 years, 343 g 38, 54 days, 7.3 cm, 2.4 g 14, 92 days and 6.2 g, 461 days and 101.7 g 43.
Observations age, TOTAL LENGTH, and weight LAB: 0+: average 8.8-11.5 cm, 6.7-20 g, 1+: average 14.6-15 cm, 30.2-36.9 g 34, 4 cm, 0.5 g 35.
Sexual maturity for at least 50% of JUVENILES: males: 1-2 years, 25.6-26.4+ cm, 424 g, females: 1.5-3 years, 26.4-32.2+ cm, 553 g:
- Observations age and TOTAL LENGTH WILD: females from 3 years, 37.8 cm fork length, 552.9 g on, males from 2 years, 35.0 cm fork length, 424.1 g on: Stanborough lake, Hertfordshire, and Ash Drain (Great Ouse Middle Level), Cambridgeshire, England (introduced) 20, male 50% maturity: 11 months, 26.4 cm, female 50% maturity: 18 months, 32.2 cm 16, females 50% maturity: 26.4 cm, males 50% maturity: 25.6 cm: Seyhan Dam lake, Turkey (introduced) 23.

Maturation and manipulation: ambient temperatures promote maturation (further research needed)

Maturation and PHOTOPERIOD manipulation: no data found yet.
Maturation and temperature manipulation:
- FARM: JUVENILES in 500 L tanks at either 6, 9, 12, 15, 23 °C. After five months, in females, higher gonad weight relative to body weight under 9-15 °C than under 6 or 23 °C (3.9-4.7% versus 0.8-1.2%). In males, higher gonad weight relative to body weight under 9 and 12 °C than under 6 or 15-23 °C (0.6-0.8% versus 0.1-0.4%) 38.

Adults: 4-14 years, 31-130 cm, 0.7-18 kg

Observations age, TOTAL LENGTH, and weight WILD: 71 cm fork length and 3.7 kg: Relief and Cut-off Channel, England (introduced) 31-4, 90 cm fork length and 7.8 kg, 13 years and mean 83 cm fork length: Fenland Waters, England 4, 8 years, 70.2 cm: Stanborough lake, Hertfordshire, England (introduced) 20, 1.3-3.9 kg: Elbe river, Czech Republic 15, 13-14 years, ca 90 cm: Castillon reservoir, France (introduced) 21, average 55 cm, 1,384 g: Aras lake dam, Azerbaijan 14, commonly 50-70 cm total length and 2-5 kg, maximum length 130 cm with a weight of 12-18 kg 44.
Observations age, TOTAL LENGTH, and weight LAB: 31-39 cm 3, 682 g 45, 4-5 years, male average 1,195 g, female average 1,662 g, average 53.7 cm 46, female average 710.8 g, male average 1,021.8 g 11, 4-5 years, average 1,367.4 g, 53.7 cm 33.

10.2 Sexual conversion

No data found yet.

10.3 Sex ratio

No data found yet.

10.4 Effects on growth

Growth rate: 51 mm/year for large juveniles (further research needed)

Natural growth rate:
- WILD: for JUVENILES of 25-30 cm, average 51 mm/year until they reached 40 cm 7.

Growth and size-grading: no effect (further research needed)

LAB: FRY in 40 dm³ tanks with unlimited access to food and either unsorted, in groups of small size, or in groups of large size. After 21 days, no difference in specific growth rates (9.2-10.1%/d) but higher survival in small-size and large-size (52.7-59.7% versus 39.0%) than unsorted groups. No difference in cannibalism rate between large-size and unsorted groups but lower rate in small-size groups (11.3% versus 21.3-29.0%). The latter is due to lower cannibalism rates in weeks 1 and 2. By week 3, no differences in cannibalism rates between three groups. Higher manipulation losses in small-size (14.7% versus 2.7%) than large-size groups; unsorted groups in between (8.0%) 42.

For growth and...
...light intensity ➝ D15,
...light colour ➝ D16,
...water temperature ➝ D17,
...salinity ➝ D18,
...stocking density ➝ D19.

10.5 Deformities and malformations

Deformities and malformations: non-inflated swim bladders (further research needed)

Observations LAB, FRY: non-inflated swim bladders 42.

11 Reproduction

11.1 Nest building

Nest building: male builds nest in sand or gravel or uses artificial substitute

Nest building and substrate:
- WILD, ADULTS: male builds nest in sand or gravel or among exposed roots in turbid waters at 1-3 m depth. Nest size typically 50 cm in diameter and 5-10 cm deep 47.
- LAB: pairs of ADULTS photo-thermally manipulated and injected with carp pituitary hormone in 600 L glass pools with synthetic material nest. Male claimed nest by surrounding it in shape of lying "8". Before and after spawning, male cleaned nest from mud and colloids to assure oxygen supply for eggs 11.
Nest building and water velocity: no data found yet.
Nest building and water depth: ➝ Nest building and substrate.
Nest building: no data found yet.

11.2 Attraction, courtship, mating

Attraction: intensified body colour, iridescent stripe, swollen opercular in males, swollen genitals in females (further research needed)

Attraction and body colour: ➝ D20.
Signals of maturity: ➝ D20.

Courtship sequence: male and female encircle nest, male body colour and marking changes (further research needed)

Courtship sequence:
- LAB: pairs of ADULTS photo-thermally manipulated and injected with carp pituitary hormone in 600 L glass pools with synthetic material nest. Male vertically shook head and circled over the nest. Cleaned nest to signal where female may place eggs. Allowed female access to nest with increasing frequency. When being chased away, female encircled nest in shape of lying "8" with upward-held head while male shook head. When pair stabilised, male body colour intensified, iridescent stripe appeared, opercula swelled, male repetitively raised dorsal fins and showed teeth. Male and female encircled nest, female with head up, sometimes in zigzag moves, male with head down. Movements changed to slow side-by-side circling above the nest for hours to days. Female's genitals swelled, she moved in mounting position. Shortly before spawning, male encircled female in downward almost vertical position, shook head quickly 11.
Courtship duration: several hours to several days ➝ Courtship sequence.

11.3 Spawning

Spawning conditions: February-May, 10-15 °C, at any time of the day

Spawning substrate: ➝ D1.
Spawning season: February-May:
- Observations WILD: spring: Curonian lagoon, Baltic Sea, Lithuania 8, mid-March to early May, 12-15 °C: Fumemorte canal, Rhone delta, southern France 16, February-March, 10.4 °C: Seyhan Dam lake, Turkey (introduced) 23.
- Observations FARM: usually May, at 12-15 °C 32.
Spawning (day)time:
- LAB: two mixed-sex pairs of ADULTS each in 50 cm deep 1,500 L tanks under lighting regimes of either "12 h light and 12 h dark", 24 h light, or 24 h dark for 40 days. Earlier spawning (within 14 days) under 24 h PHOTOPERIOD than other regimes. Spawning at all times of the day. No interaction between PHOTOPERIOD and spawning quality in terms of nest coverage but no single good quality (i.e. 100% coverage) spawning in 24 h dark condition. No difference in cortisol levels 46.
- LAB: pairs of ADULTS photo-thermally manipulated and injected with carp pituitary hormone in 600 L glass pools with synthetic material nest spawned at any time of the day 11.
Spawning temperature: no data found yet.
Spawning salinity: no data found yet.
Spawning and water velocity: no data found yet.
Spawning depth: no data found yet.
Spawning density: no data found yet.

Male:female ratio resulting in spawning, composition of the broodstock: 1:1 (further research needed)

Male:female ratio resulting in spawning:
- Observations LAB, ADULTS: 1:1 46.
Composition of broodstock: no data found yet.

Spawning sequence: female spawns close to nest, male 7 s later during 1-3 sessions (further research needed)

Spawning sequence:
- LAB: pairs of ADULTS photo-thermally manipulated and injected with carp pituitary hormone in 600 L glass pools with synthetic material nest. For spawning, female moved swollen light pink genitalia close to nest, bend caudal fin, and released eggs in average 2 s. Male behind her almost motionless. Female left nest quickly. Average 7 s later, male released milt over eggs by lateral winding in average 19 s during 1-3 sessions 11.
Spawning duration: ➝ Spawning sequence.

11.4 Fecundity

Female fecundity: average 140,000-255,000 eggs/kg body weight

Number of spawns: no data found yet.
Fecundity per spawn:
- Observations absolute fecundity: no data found yet.
- LAB, ADULTS: females of average 710.8 g had lost 54.8 g weight after spawning 11.
- Observations relative fecundity WILD, ADULTS: average 140,000-225,000 eggs/kg body weight: Stanborough lake, Hertfordshire, and Ash Drain (Great Ouse Middle Level), Cambridgeshire, England (introduced) 20, mean 255,412 eggs/kg: Fumemorte canal, Rhone delta, southern France 16, 170,000-230,000 eggs/kg body weight 44.

Male fecundity: 1-3 times per spawning (further research needed)

Number of spawns: ➝ D21.
Fecundity per spawn:
- Observations absolute fecundity: no data found yet.
- Observations relative fecundity: no data found yet.
- LAB, ADULTS: males of average 1,021.8 g had lost 52.5 g weight after spawning 11.

11.5 Brood care, breeding

Breeding type: gravel breeder

For nest building ➝ D1.

12 Senses

12.1 Vision

Visible spectrum: green, red (further research needed)

LAB: retina of dissected eyes of JUVENILES absorbed wavelengths of 400-700 nm with peaks at 535 nm (green) and 603 nm (red) 36.

Importance of vision: hunting (further research needed)

Vision and hunting:
- WILD, ADULTS: visual hunter: more diurnal movement at new moon (ca 150 m versus 60-100 m) compared to all other moon phases, probably due to lower predation success in dark nights: Elbe river, Czech Republic 15.

12.2 Olfaction (and taste, if present)

No data found yet.

12.3 Hearing

No data found yet.

12.4 Touch, mechanical sensing

No data found yet.

12.5 Lateral line

No data found yet.

12.6 Electrical sensing

No data found yet.

12.7 Nociception, pain sensing

No data found yet.

12.8 Other

No data found yet.

13 Communication

13.1 Visual

No data found yet.

13.2 Chemical

No data found yet.

13.3 Acoustic

No data found yet.

13.4 Mechanical

No data found yet.

13.5 Electrical

No data found yet.

13.6 Other

No data found yet.

14 Social behaviour

14.1 Spatial organisation

Aggregation type: no school (further research needed)

WILD: no schooling behaviour: river Oulujoki, Finland 9.

Stocking density and stress: direct relation from >50 ind/L on (further research needed)

FARM: FRY in 200 L cylindro-conical rearing tanks under 24 h PHOTOPERIOD. In first experiment under densities of 25, 50, or 100 IND/L. After 14 days, higher survival under low and middle than under high density (78.5-79.2% versus 72.3%).
In subsequent second experiment, 18 day-old FRY from first experiment mixed and objected to densities of 6, 10, or 15 IND/L. After 21 days, higher survival under low than under middle or high density (56.5% versus 45.4-48.4%). Lower type II cannibalism under low than under high density (27.2 versus 35.2%); middle density in between (32.4%) 18.

Stocking density and growth: inverse relation from >50 ind/L on (further research needed)

FARM: FRY in 200 L cylindro-conical rearing tanks under 24 h PHOTOPERIOD. In first experiment under densities of 25, 50, or 100 IND/L. After 14 days, no difference in swim bladder inflation (90.1-90.5%), but higher weight and higher length under low than under high density (38.8 versus 27.8 mg, 16.1 versus 14.6 mm); middle density in between (34.3 mg, 15.5 mm). Daily growth rate higher under low than under high density (2.7 versus 1.9 mg/d, 0.8 versus 0.6 mm/d); middle density in between (2.4 mg/d, 0.7 mm/d).
In subsequent second experiment, 18 day-old FRY from first experiment mixed and objected to densities of 6, 10, or 15 IND/L. After 21 days, higher weight and higher length under low than under high density (640.0 versus 520.0 mg, 44.2 versus 41.5 mm); middle density in between (610.0 mg, 43.8 mm). Daily growth rate higher under low than under high density (28.8 versus 23.1 mg/d, 1.4 versus 1.2 mm); middle density in between (27.4 mg/d, 1.3 mm/d) 18.

14.2 Social organisation

No data found yet.

14.3 Exploitation

Cannibalism, predation: prevalent

For cannibalism and...
...feeding ➝ D3,
...size-grading ➝ D22,
...stocking density ➝ D23.

14.4 Facilitation

Cooperation, mutualism: attacks in packs (further research needed)

WILD: packs of ADULTS attacked larger prey which died 2-4 h later: Great Ouse Relief Channel, England (introduced) 4.

14.5 Aggression

No data found yet.

14.6 Territoriality

No data found yet.

15 Cognitive abilities

15.1 Learning

No data found yet.

15.2 Memory

No data found yet.

15.3 Problem solving, creativity, planning, intelligence

No data found yet.

15.4 Other

No data found yet.

16 Personality, coping styles

No data found yet.

17 Emotion-like states

17.1 Joy

No data found yet.

17.2 Relaxation

No data found yet.

17.3 Sadness

No data found yet.

17.4 Fear

No data found yet.

18 Self-concept, self-recognition

No data found yet.

19 Reactions to husbandry

19.1 Stereotypical and vacuum activities

No data found yet.

19.2 Acute stress

Handling: collection and anaesthesia is stressful (further research needed)

Collection and anaesthesia:
- LAB: most FRY of 4-18 days post hatch collected for length and weight measurements and anaesthetised in 0.01-0.05% Propiscin solution did not survive procedure 18.

Confinement: stressful if done for 1 h (further research needed)

LAB: single ADULTS per tank were netted, confined to half-full 30 L tank for 1 h, and returned to rearing tank. After 48 h, higher cortisol levels in handled and confined compared to control ADULTS (59.0 versus 40.8 ng/mL) indicating stress 19.

Live transport: stressful (further research needed)

WILD: trawler bycatch of undersized individuals of <370 mm were either emptied into 500 L tank containing lake water of 15-21.4 °C ("no chilling" group) or ice water of 0 °C ("10 min chilling" group). After 10 min it took to separate undersized individuals from catch, they were transferred to 50 L transport tank containing lake water. Third group was left in 500 L ice water tank for 120 min ("120 min chilling" group).
a) Mortality: directly after treatment, higher mortality under 120 min chilling than under 10 min chilling and no chilling (73.9% versus 2.6-6%). Higher mortality the smaller the TOTAL LENGTH, with mortality ca 90% at 100-150 mm length. After 48 h recovery period, higher mortality under 120 min chilling than under 10 min chilling or no chilling (66.7% versus 22.6-26.3%). Same effect of body size. Overall mortality: 91.3% after 120 min chilling, 27.2% after 10 min chilling, 28.2% after no chilling.
b) Stress: directly after treatment, higher cortisol levels in trawled and handled individuals than control individuals (ca 180-380 ng/mL versus ca 75 ng/mL). Peak of cortisol levels in all treatments around 20-40 min afterwards. After 48 h recovery, higher cortisol levels than control (ca 200-300 ng/mL versus ca 75 ng/mL) 13.
FARM: JUVENILES stocked in earthen ponds at 350,000 IND/ha. Directed from pond through channel to loading basin and transported at 7,500 IND/m³. Higher cortisol levels in loading basin than in pond and channel (622.3 ng/mL versus 262.7 ng/mL). Back to pre-stress levels at 1.5 and 24 h after transport (333.5 ng/mL). Relatively high cortisol levels pre- and post-stress indicate stress through density and handling. Higher glucose levels 1.5 h after transport than in pond and channel (195.3 mg/dL versus 105.3-123 mg/dL); loading basin in between. Back to pre-transport values at 24 h after transport (93 mg/dL) 14.

19.3 Chronic stress

Handling: delays or prevents spawning despite adaptation to repeated stress (further research needed)

LAB: two mixed-sex pairs of ADULTS each in 50 cm deep 1,500 L tanks under lighting regimes of either "12 h light and 12 h dark", 24 h light, or 24 h dark. Half of the pairs were stressed by confining ADULTS to 10 cm water level, capturing them, and holding them outside the water for 20 s each day. During 40 days, 29 of 36 females spawned in artificial nests, seven non-spawners from stress groups. Delayed spawning in stressed compared to unstressed groups across all PHOTOPERIODs. No difference in cortisol levels indicating adaptation to long-term stress 46.

For chronic stress and...
...PHOTOPERIOD ➝ D24,
...stocking density ➝ D23.

19.4 Stunning reactions

Stunning rules: fast, effective, safe

Stunning rules: to minimise pain reactions and enhance welfare before slaughter:
1. induce insensibility as fast as possible,
2. prevent recovery from stunning,
3. monitor effectiveness (observations, neurophysiological measurements) 48.

Stunning methods: electrical stunning most effective (further research needed)

Electrical stunning:
- LAB: ADULTS restrained in unit of 150 x 30 x 40 cm with 18 rows of 14 electrodes each connected to 100 Hz AC/DC electrical current. Combination of 144 V dc and 13 Vrms ac 100 Hz through the head for
  a) 1 s yielded general epileptiform insult in 25 of 25 ADULTS for 23 s. Of all ADULTS, one displayed behaviour, 15 brain activity to needle scratches 30 s after the stun. 22 of 25 ADULTS had 15 s heart fibrillation after the stun. Then, heart rate increased compared to pre-stun values for the first minute after the stun and returned to pre-stun values.
  b) 5 s. After stun placed in ice water. General epileptiform insult in 12 of 12 ADULTS for 115 s. No displayed behaviour to needle scratches until 5 min after stun but brain activity in one individual after 2 min. 12 of 12 ADULTS had 11 s heart fibrillation after the stun. Then, heart rate decreased compared to pre-stun values and never returned to pre-stun values 45.

Glossary

ADULTS = mature individuals, for details ➝ Findings 10.1 Ontogenetic development
FARM = setting in farming environment or under conditions simulating farming environment in terms of size of facility or number of individuals
FOOD CONVERSION RATIO = (food offered / weight gained)
FRY = larvae from external feeding on, for details ➝ Findings 10.1 Ontogenetic development
IND = individuals
JUVENILES = fully developed but immature individuals, for details ➝ Findings 10.1 Ontogenetic development
LAB = setting in laboratory environment
MILLIARD = 1,000,000,000 29 30
PHOTOPERIOD = duration of daylight
TOTAL LENGTH = from snout to tip of caudal fin as compared to fork length (which measures from snout to fork of caudal fin) 41 or standard length (from head to base of tail fin) or body length (from the base of the eye notch to the posterior end of the telson)
WILD = setting in the wild

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