Patterns of the electric organ discharge during courtship and spawning in the mormyrid fish,Pollimyrus isidori

Summary Pollimyrus isidori's electric organ discharge (EOD) is of the pulse type. Patterns of EOD intervals were investigated prior to, during and following spawning behaviors as related with overt behaviors, and with the sound production by the nestbuilding male. Prior to the time of reproduction, isolated and socially interacting fish (n=15) showed characteristic discharge interval patterns for resting, swimming, probing, hovering and hiding activities. Males (n=8) and females (n=6) did not differ in their mean EOD repetition rates during resting (11.6±2.5 Hz), nor Short Bursts/min (less than 20 intervals of 8–13 ms). In interacting fish Long Bursts (greater than 20 intervals of 8–13 ms, lasting for more than 300 ms) were observed only during the attack and bite sequence. A pursuing fish displayed a rapid alternation of Long Bursts with Discharge Breaks (300–1000 ms silence) during the chase behavior. Avoidance behavior which followed from several attacks was correlated with a Medium Uniform Rate (8–12 Hz) normally lasting for 20 to 60 s, or a Discharge Arrest (silence greater than 1 s) in the submissive fish. The nocturnal courtship behavior began soon after dark (1900 h). Spawning typically started 2 to 5 h after dark, continuing for 2 to 6 h until about 0200 h. During courtship and spawning the female's brief visits (15–25 s) to the male's territory recurred every 30–60 s. At all other times the female was aggressively excluded from the nest region. Courtship and spawning behaviors are described along with the electrical displays identified from 19 spawnings in three fish pairs (from a total of 37 spawnings in 4 males and 4 females). Just prior to the onset of courtship behavior, with male territorial aggression beginning to decline, females switched from a Medium Sporadic Rate pattern (resting and hiding patterns; 13 Hz) to a Medium Uniform Rate pattern (6–8 Hz) while still in their hiding area. Females continued to display this uniform rate throughout the courship and spawning period, including the courtship and spawning bouts when Discharge Breaks or Arrests also occurred. This persistance distinguishes the courtship pattern from the similar avoidance pattern (see above). The male courtship and spawning EOD pattern was similar to the female's and unique for a territorial male. He switched from a High Sporadic Rate (swimming EOD pattern; about 18 Hz) to a regularized Medium Uniform Rate (about 9 to 11 Hz) only during courtship and spawning bouts, including 1–3 EOD Breaks during Vent-to-Vent coupling (average interval: 272±71 ms, n=37). No sooner had the female left the spawning site than he resumed displaying a High Sporadic Rate. This temporal correlation of reproductive behaviors with electrical displays suggests their instrumental role in mutual acceptance of mates. Males showed their sex-specific type of EOD phase-locking, the Preferred Latency Response, only during the first few hours of entry of a fish in their tank. Two females with EOD waveform features more typical of males also spawned repeatedly; waveform does not appear to be critical. Males stopped their nocturnal sound production for the later part of courtship and the whole spawning period. Except for infrequent attacks on the female between spawning bouts, the male did not resume singing until the end of spawning when all eggs were shed (around 0200 h); from this time on the male sang until dawn. The sequencing of the three acoustic elements (moans, grunts, growls) are described. A catalogue of discharge patterns correlated with overt behaviors (Tables 1, 2), and an integrated summary time table of P. isidori's complex reproductive behavior are presented.     

Electrosensory prey detection in the African sharptooth catfish, Clarias gariepinus (Clariidae), of a weakly electric mormyrid fish, the bulldog (Marcusenius macrolepidotus)

During their seasonal runs in the Okavango and other freshwater bodies in southern Africa, the sharptooth catfish, Clarias gariepinus, specialises on the bulldog, Marcusenius macrolepidotus, as its main prey. We examined whether the catfish can locate bulldogs by detecting their pulse-type electric organ discharges (EODs). The electrosensory threshold for single-cycle, monopolar square-wave pulses was exceedingly low (down to 13 μV_p–p/cm for 4-ms pulses) in trained, food-rewarded sharptooth catfish (n=4), confirming the results of Lissmann and Machin who were, however, unable to identify a biological function. Other stimulus pulse waveforms (single-cycle, monopolar as well as bipolar sine-wave pulses) were also effective stimuli according to their spectral low-frequency energy contents (0– 30 Hz). Male bulldogs display an EOD pulse approximately 10× the duration of female EODs (≈0.5 ms). The C. gariepinus threshold for field-recorded playbacks of a male bulldog EOD (of long duration) was 103 μV_p–p/cm, whereas the brief female and juvenile EODs were not detected (using intensities of natural EODs). EODs of other mormyrids were detected when either monopolar or of long duration. Signal source amplitude increased linearly with standard length (SL) in bulldogs. Signal reach, as calculated from signal source amplitude and receiver sensitivity, is up to 150 cm for a large male bulldog (SL 27.5 cm), and 83 cm for a male that has just turned sexually mature (SL 12.6 cm). Therefore, most bulldogs eaten by catfish are probably male, in agreement with the size distribution of bulldogs found in catfish stomachs. These results suggest that sharptooth catfish rely heavily on their acute electrical sense during hunting, and an important function for electroreception in an African catfish has been identified. Received: 13 December 1999 / Revised: 20 April 2000 / Accepted: 20 May 2000     

Strategic male signalling effort in a desert-dwelling fish

Males often use elaborate courtship displays to attract females for mating. Much attention, in this regard, has been focused on trying to understand the causes and consequences of signal variation among males. Far less, by contrast, is known about within-individual variation in signal expression and, in particular, the extent to which males may be able to strategically adjust their signalling output to try to maximise their reproductive returns. Here, we experimentally investigated male courtship effort in a fish, the Australian desert goby, Chlamydogobius eremius. When offered a simultaneous choice between a large and a small female, male gobies spent significantly more time associating with, and courting, the former, probably because larger females are also more fecund. Male signalling patterns were also investigated under a sequential choice scenario, with females presented one at a time. When first offered a female, male courtship was not affected by female size. However, males adjusted their courtship effort towards a second female depending on the size of the female encountered previously. In particular, males that were first offered a large female significantly reduced their courtship effort when presented with a subsequent, smaller, female. Our findings suggest that males may be able to respond adaptively to differences in female quality, and strategically adjust their signalling effort accordingly.     

The ecology,discharge diversity and predatory behaviour of gymnotiforme electric fish in the coastal streams of French Guiana

Summary 1. A survey of the species distribution, discharge characteristics, ecology and behaviour of South American Gymnotiforme electric fish was carried out during two field trips to sites in the vicinity of Kourou, French Guiana, in 1983 and 1985. 2. Measurements of water conditions, collected over a twelve month period, highlight the strong ecological pressure on species exploiting the rapidly fluctuating and often temporary coastal streams. Adaptation of the reproductive cycle to the rainy seasons is particularly marked and the possible zeitgebers are considered. Given the almost invariant water conductivity, its suggested use as an environmental trigger is rejected in favour of acoustic or mechanical cues. 3. The two main study areas were inhabited principally by pulse gymnotoids of the genera Gymnotus and Hypopomus. A new Hypopomus species was discovered as well as specimens of G. pantherinus, previously undescribed in French Guiana. Larvae of H. beevei were found prior to the start of the 1985 rainy season. Developmental data for this species are presented which support the view that, as in other weakly electric gymnotoids, a primitive larval electric organ probably precedes the adult structure. The larval discharge also strongly resembles that of the adult electric eel, lending weight to published theories of an Electrophorus-like ancestor to the wekly electric fish. 4. The predatory behaviour of the strongly electric eel was recorded in detail. The field results indicated that this species preyed on weakly electric fish whose responses to the foraging eel are also reported. The probable use of electrical cues by hunting eels was further investigated in a laboratory experiment in which predatory responses were compared in the presence of non-electric and electric live prey, and models mimicking electric prey. It is concluded that electrical cues are probably of paramount importance in the eel's normal prey capture behaviour.     

Nest distribution affects behaviour and mating success in a marine fish

The distribution of breeding resources, such as nest sites, can have a pronounced impact on a population by affecting the proportion of individuals that succeed to breed and hence, the variation in reproductive success. Aggregation of important resources can lead to resource monopolisation by a limited number of individuals and thus affect the intensity of sexual selection. In this study, we tested, by contrasting two experimental treatments (dispersed vs. aggregated), how nest distribution affects: (1) mating behaviour, (2) male nest occupation and mating success, and (3) reproductive success and the opportunity for selection. We used the two-spotted goby (Gobiusculus flavescens), a small marine fish with a resource-based mating system, as our model species. When nests were aggregated, a larger proportion of the males behaved aggressively, fewer males succeeded in occupying a nest, fewer males became mated, and those males that mated received fewer eggs from spawning females. These effects resulted in a higher variance in reproductive success and hence, a higher opportunity for selection (I _rs ), in the aggregated treatment. We suggest that the results are a direct consequence of males defending a territory around their nest, preventing competitively inferior males from breeding. However, we found no significant selection differentials for body length or condition of males in either treatment. Our results support the hypothesis that aggregation of essential resources like nests promotes resource monopolisation. In species facing highly clumped nesting resources in the wild, monopolisation may negatively impact population productivity but could lead to strong selection on traits that promote male competitive ability.     

Spawning behaviour,egg and larval development of the myctophid fish Benthosema pterotum

The distribution of eggs and early larval stages of the myctophid fish Benthosema pterotum was studied during three cruises to the Gulf of Oman during January to February 1981 and February 1983. The eggs and yolk sac larvae were identified and described after artificial fertilization on board the Research Vessel Dr. Fridtjof Nansen. The mature fish spawn in the Gulf at 300 to 100 m depths during early night. The eggs and smallest larvae were found at 300 to 200 m depths. The eggs hatch within 12 h at 21°C. Hatching occurs before the slightly bouyant eggs reach the upper 50 m of the water column.     

Predicting fish behaviour in response to abyssal food falls

Estimates of fish abundance based on arrival rates and numbers present at baited cameras allow multiple, replicate assessments where data are not available by other means. Unfortunately such estimates are strongly affected by the assumed behaviour of the fish species concerned. Three of the possible foraging strategies of deep-sea fish were modelled and the likely patterns of fish arrival calculated for the same fish density, swimming and current velocities and odour plume properties. Cross-current foraging resulted in the highest numbers of fish at bait, with arrival rates that fitted well to field data. The sit-and-wait strategy produced lower arrival rates with passive drifting animals arriving slowest. Each model produces a distinctive pattern of animal arrivals that may be diagnostic of each foraging strategy. The advantages, disadvantages and likely metabolic and sensory demands of each strategy are discussed. Published online: 17 September 2002     

Mating and its effect on acoustic signalling behaviour in a primitive orthopteran,Cyphoderris strepitans (Haglidae): the cost of feeding females

Summary Males of the primitive orthopteran, Cyphoderris strepitans, provide their mates with two types of nuptial food gift during mating: 1) females feed while coupled on the fleshy metathoracic wings of the male and the resultant flow of haemolymph and 2) the spermatophore transferred by the male includes a gelatinous spermatophylax which the female eats after mating. During the peak breeding interval, virgin males secure significantly more matings than their numbers relative to non-virgin males would predict. We tested the hypothesis that non-virgin males, having lost a substantial portion of available energy through previous investment in females, call significantly less than virgin males. Reduced calling should result in the attraction of fewer females and/or a greater risk of intrusion from competing males and consequently, a lowered mating success. Calling activity of male C. strepitans of varying mating status was monitored with a sound activated relay apparatus for two consecutive nights following their capture. Males were of three experimental groups, virgin, freshly wounded males (mated on the same night of capture), and old-wound males (mated at least one night prior to capture). Our cata showed a significant short-term reduction in signalling activity as a consequence of mating. Whereas there was no difference in the time spent calling by virgins on the first and second nights following capture, freshly wounded and oldwound males called significantly less on the first night than on the second. Furthermore, the proportion of males calling on both nights was significantly greater for the virgin and old-wound groups than for freshly wounded males. When the duration over which males called on the first night was compared with that of the second, virgin males showed no difference whereas old-wound males called for significantly shorter intervals on the first night. These results indicate that the cost of a large nutrient investment in females lowers the energy level in males below the threshold required for a prolonged signalling period. After a refractory interval during which males feed and replenish their energy reserves, calling levels equivalent to those which occur prior to mating are regained. Active female choice may also contribute to the reduced mating success of non-virgins, but this possibility remains untested.     

Female discharges are more electrifying: spontaneous preference in the electric fish,Eigenmannia (Gymnotiformes,Teleostei)

Summary The tropical South American teleost Eigenmannia lineata showed a spontaneous preference for the female type, compared with the male type, of its sexually dimorphic, weak-electric organ discharge (EOD). Female and male EODs differ in waveform and harmonic content. An isolated fish was simultaneously stimulated with digitally synthesized natural male and female EODs of equal peak-to-peak amplitudes, at ±35 Hz frequency difference centered on its stable resting discharge frequency. The stimulus dipoles were arranged symmetrically to the right and left of the fish's hiding place. All stimulus conditions were permuted at random sequence. Among 11 fish tested, 8 showed a statistically significant preference for one stimulus, the female type, as measured by the amount of time a fish spent close to a stimulus dipole (P<0.05 in each fish, two-tailed). Thus female EODs rather than male EODs were more attractive to adult and juvenile fish of both sexes. It was also concluded that E. lineata is capable of discriminating female from male EODs by a complex sensory capacity requiring neither amplitude nor frequency cues. The EOD waveform changed very little within the ecological range of water conductivities (approximately 10–100 S·cm^-1); the P/N-ratio (a waveform character based on zerocrossing intervals) depended only weakly, but significantly, on conductivity (negative correlation in all four fish). Also, the effect of temperature on EOD waveform was very weak: Q ₁₀-values of the P/N-ratio were below but close to 1 in all fish (27±5°C). Thus, it can be concluded that the EOD waveform is remarkably stable within widely changing conditions-even beyond the variation found in the field-and is therefore potentially useful as a social cue.     

Past reproductive effort affects parental behaviour in a cichlid fish,Cichlasoma nigrofasciatum: a comparison of inexperienced and experienced breeders with normal and experimentally reduced broods

Defensive and parental care behaviour of convict cichlids that differed in past effort was compared. Before testing, some fish were bred three times while others were not bred. Age was held constant; all individuals in this study were approximately 20 months old (±2 months) at test time. Furthermore, half of the pairs in this study had their broods experimentally reduced by 50%. Results indicated that past effort across breeding attempts affects investment in the current brood. Experienced pairs were more aggressive toward a model predator than inexperienced parents. However, no major differences were observed in depreciable care (i.e. fanning). Contrary to previous studies, brood size had minor effects on parental care. This discrepancy could be due to the age of the parents; individuals in this study were significantly older than fish tested in previous studies. The results support parental investment theory and suggest that past effort is not only important within breeding episodes but also within an animal's lifetime.