# These authors contributed equally 6 *Corresponding author: Stefan Mucha -stefan.mucha@hu-berlin.de 7 Keywords: dissolved oxygen, swimming behaviour, active sensing, electric organ discharge, gymnotiform, 8 shuttle-box choice chamber 9 Summary 10The weakly electric knifefish, Apteronotus albifrons, avoids hypoxia below 22% air saturation. Avoidance 11 correlates with increased swimming activity, but not with a change in electric organ discharge frequency. 12 Abstract 13Anthropogenic environmental degradation has led to an increase in the frequency and prevalence of 14 aquatic hypoxia (low dissolved-oxygen concentration, DO), which may affect habitat quality for water-15 breathing fishes. The weakly electric black ghost knifefish, Apteronotus albifrons, is typically found in 16 well-oxygenated freshwater habitats in South America. Using a shuttle-box design, we exposed juvenile 17A. albifrons to a stepwise decline in DO from normoxia (>95% air saturation) to extreme hypoxia (10% air 18 saturation) in one compartment and chronic normoxia in the other. Below 22% air saturation, A. 19 albifrons actively avoided the hypoxic compartment. Hypoxia avoidance was correlated with upregulated 20 swimming activity. Following avoidance, fish regularly ventured back briefly into deep hypoxia. Hypoxia 21 did not affect the frequency of their electric organ discharges. Our results show that A. albifrons is able 22 to sense hypoxia at non-lethal levels and uses active avoidance to mitigate its adverse effects. 23

To understand animal ecology, observation of wildlife in the natural habitat is essential, but particularly challenging in the underwater realm. Weakly electric fishes provide an excellent opportunity to overcome some of these challenges because they generate electric organ discharges (EODs) to sense their environment and to communicate, which can be detected non-invasively. We tracked the EOD and swimming activity of two species of mormyrid weakly electric fishes (Marcusenius victoriae and Petrocephalus degeni) over diel cycles in the laboratory, and we recorded EODs and environmental dissolved oxygen (DO) concentration and temperature over several months in a naturally hypoxic habitat in Uganda. Under laboratory conditions, both species showed increases of activity and exploration behavior that were closely synchronized to the onset of the dark phase. In the wild, fish preferred structurally complex habitats during the day, but dispersed toward open areas at night, presumably to forage and interact. Nocturnal increase of movement range coincided with diel declines in DO concentration to extremely low levels. The fact that fish showed pronounced nocturnal activity patterns in the laboratory and in the open areas of their habitat, but not under floating vegetation, indicates that light intensity exerts a direct effect on their activity. We hypothesize that being dark-active and tolerant to hypoxia increases the resistance of these fish against predators. This study establishes a new technology to record EODs in the field and provides a window into the largely unknown behavior of mormyrids in their natural habitat.

Anthropogenic environmental degradation has led to an increase in the frequency and prevalence of aquatic hypoxia (low dissolved oxygen concentration, DO), which may affect habitat quality for water-breathing fishes. The weakly electric black ghost knifefish, Apteronotus albifrons, is typically found in well-oxygenated freshwater habitats in South America. Using a shuttle-box design, we exposed juvenile A. albifrons to a stepwise decline in DO from normoxia (> 95% air saturation) to extreme hypoxia (10% air saturation) in one compartment and chronic normoxia in the other. On average, A. albifrons actively avoided the hypoxic compartment below 22% air saturation. Hypoxia avoidance was correlated with upregulated swimming activity. Following avoidance, fish regularly ventured back briefly into deep hypoxia. Hypoxia did not affect the frequency of their electric organ discharges. Our results show that A. albifrons is able to sense hypoxia at non-lethal levels and uses active avoidance to mitigate its adverse effects.

Hypoxia is a widespread environmental stressor that shapes fish physiology and morphology. Plasticity in traits that improve oxygen uptake and delivery or reduce oxygen requirements may be critical for fish to cope with fluctuating dissolved oxygen (DO) conditions in their natural habitat or adapt to new environments. In this study, we characterized a suite of morpho-physiological respiratory traits of a naturally hypoxia-acclimated weakly electric mormyrid fish, Petrocephalus degeni, and quantified their plasticity in response to long-term normoxia exposure. We captured P. degeni from a hypoxic swamp habitat (PO2 = 2.43 ± 1.85 kPa) surrounding Lake Nabugabo, Uganda, and acclimated them to normoxia (PO2 > 16 kPa) for up to 75 days. At various time points throughout normoxia exposure, we measured blood hemoglobin and lactate concentration, gill size, routine metabolic rate (RMR), regulation index (RI), and critical oxygen tension (Pcrit). We found that 62–75 days of normoxia exposure significantly reduced blood hemoglobin concentration (− 17%), gill filament length (− 14%), and hemibranch area (− 18%), whereas RMR, RI, Pcrit, and blood lactate showed no significant change. Our results support earlier findings that swamp-dwelling P. degeni are well adapted to life in chronic and severe hypoxia and indicate that they possess a limited capacity for phenotypic plasticity in response to a change in their DO environment.