Electric communication during courtship and spawning in two sibling species of dwarf stonebasher from southern Africa, Pollimyrus castelnaui and P. marianne (Mormyridae, Teleostei): evidence for a non species-specific communication code?

The fixed part of the electrocommunication signal, the electric organ discharge (EOD) waveform, is well differentiated in the two vicariant dwarf stonebasher species, Pollimyrus castelnaui and P. marianne. However, differentiation regarding the variable, situation-dependent part, i.e., inter-discharge interval (IDI) patterns, has never been studied in a pair of sibling species of mormyrid fish. We here compare the electrical signalling that accompanies different motor behaviours (such as resting and swimming, territorial agonistic interactions, courtship and spawning) in the two species. Double pulse patterns of regularly alternating short IDIs of 8-11 ms and long ones of 16-100 ms accompanied threat displays in both species. In three pairs of P. marianne and five pairs of P. castelnaui, courtship was characterised by nest building, territory patrolling and acoustic displays (advertisment calls) that were accompanied by long discharge breaks in the male and highly regular IDIs around 50 ms in the female of both species. Nest-tending males showed IDI sequences consisting of regularly alternating double pulse patterns, similar to threat displays. During spawning both sexes generated stereotyped IDI sequences of a low discharge rate. All IDI patterns occurring in one species were also found in the other, and no species-specifity was identified at that level. Playback experiments contrasting conspecific and heterospecific IDI sequences (that had been recorded from nocturnally swimming fish) revealed preferences in none of the six experimental subjects. Double pulse patterns, high discharge rate displays (HD) and regularisations of the IDI sequence accompanying specific behaviours occurred in similar form in both dwarf stonebasher species of the present study. Therefore, we conclude that in the speciation of P. castelnaui and P. marianne the fixed part of the EOD, its waveform, was under more differential selection pressure than its variable part, the patterns of IDI

Electrosensory Frequency and Intensity Discrimination in the Wave-Type Electric Fish Eigenmannia

Eigenmannia's frequency and intensity discrimination thresholds were determined for a range of seven frequencies (50-1700Hz), centered on a fish's individual discharge frequency, using a conditioned go/no-go paradigm. The threshold criterion was '50% correct choices' (of the rewarded stimulus, S+, over the unrewarded stimulus, S—); this was validated by testing for statistically significantly shorter response latencies for the S+ compared with the S— stimulus. The stimuli consisted of sine wave bursts presented for up to 20 s, repeated at 2 s"1 (rise and fall times of an individual burst, 50 ms; holding time, 250 ms; silence, 150ms). When testing for frequency discrimination, the sine wave bursts alternated in frequency (A/); for intensity discrimination, every other sine wave burst was of increased intensity (A/). The reference stimulus intensity was 30 dB with reference to a fish's individual absolute threshold for a continuous sine wave at that frequency, previously determined using a conditioned go response. Sensory discrimination was best close to a fish's individual discharge frequency. At 30 dB sensation level, fish discriminated frequency differences as small as 0.52 Hz (0.60 and 0.79 Hz in two other individuals) and intensity differences as small as 0.56dB (1 dB in two other fish). At stimulus frequencies different from a fish's discharge frequency, Eigenmannia's frequency discrimination declined at lower frequencies at a rate of up to 1 Hz octave"1, and at higher frequencies at a rate of up to 3 Hz octave"1. For Eigenmannia's intensity discrimination a similar loss was observed: at frequencies lower than a fish's discharge frequency, intensity discrimination thresholds rose at a rate of less than 1 dB octave" *, while the rate was below 2 dB octave"1 for higher frequencies. Compared with other acoustico-lateral senses in lower vertebrates, Eigenmannia's electrosensory frequency and intensity discrimination is unusually high, in the range of that known for audition in the most sensitive higher vertebrates with a cochlea (for example, human). This emphasizes Eigenmannia's specialized 'active' electrosensory system, which detects the presence of a stimulus field as the modulation of a fish's own 'carrier' signal in amplitude and phase (beat analysis), as opposed to 'passive' sensory systems, which must deal with unpredictable signals from the environment as they occur

Species recognition by the sequence of discharge intervals in weakly electric fishes of the genus Campylomormyrus (Mormyridae, Teleostei)

In two Campylomormyrus species, tamandua and rhynchophorus from Central Africa, the electric organ discharge (EOD) activity was studied during the nocturnal activity phase in the laboratory. Both species have a pulse-type EOD of less than 200 μs duration and similar waveform; the sequence of inter-discharge intervals, however, differed characteristically: C. tamandua displayed a 'micro-pattern' encompassing nearly the whole range of intervals in a sequence of only three intervals (long-intermediate-short), while in C. rhynchophorus adjacent intervals tended to be more similar in duration. Four C. rhynchophorus and five C. tamandua were tested for their preference of conspecific pulse sequences rather than those of the other species, respectively, in playback experiments. These were performed in a T-maze paradigm such that two pulse patterns, one from each species, were simultaneously presented via two electric fish dipole models. Two pulse sequences recorded from different individuals were used to represent each species; the natural EOD waveform was replaced by a square-wave pulse of approximately natural duration. The pattern pairings and the sequence of stimulus patterns followed a randomized blocks design. Each of the four C. rhynchophorus had significantly higher preference scores for the conspecific pulse patterns than for those of C. tamandua, while in C. tamandua no such discrimination was observed. It is suggested that the juvenile C. tamandua used would probably have joined mixed-species schools as reported to exist in the wild, while this appears unlikely in the more mature C. rhynchophorus. One of the functions of the inter-discharge interval code of communication in mormyrids is species recognition although this may be seen only in sufficiently mature fish

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

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

The Victoria Falls, a species boundary for the Zambezi Parrotfish, Cyphomyrus discorhynchus (Peters, 1852), and the resurrection of Cyphomyrus cubangoensis (Pellegrin, 1936) (Mormyridae: Teleostei)

Cyphomyrus discorhynchus occurs in the Zambezi River and in the linked systems of the Kwando and the Okavango. We collected specimens from both above and below the Victoria Falls and recorded Electric Organ Discharges. We found a marked degree of anatomical differentiation among the specimens from the Zambezi delta to the Victoria Falls that represents a subspecific, geographical cline, reflecting the great length (1400 km) and high ecological diversity of that river section (Lower and Middle Zambezi). We confirm that the populations above the Falls (i.e. Upper Zambezi, Kwando and Okavango) are differentiated from those below, possessing fewer dorsal fin rays (a median of 30–31 rather than 33–34). The waveforms of the electric organ discharge pulses have four phases and show geographic variation but were briefer for specimens from below the Falls. We resurrect Cyphomyrus cubangoensis (Pellegrin, 1936) for the Upper Zambezi/Kwando/Okavango system

Hippopotamyrus ansorgii species complex in the Upper Zambezi River System with a description of a new species, H. szaboi (Mormyridae)

Specimens referable to Hippopotamyrus ansorgii sampled from the Upper Zambezi River System within Caprivi (Namibia) represent a complex of three species, two of which coexist in the Upper Zambezi River, and a third that inhabits a nearby river, the Kwando, with which the Zambezi has been connected during periods of flooding. All three are indistinguishable in terms of their general appearance, but differ consistently in electric organ discharges (EOD), morphology, and molecular genetic characters. All phenotypes display a monopolar, headpositive EOD pulse with specific post- or prepotentials. For H. ansorgii from the Zambezi River (HaZ), pulse duration is less than 0.5 ms (down to 0.205 ms; N = 34); for the syntopic H. szaboi sp. n., it is greater than 0.6 ms (up to 1.8 ms at 10% peak amplitude; N = 19). The parapatric phenotype of H. ansorgii from the Kwando River (HaK) has pulses shorter than 0.215 ms (down to 0.105 ms; N = 36). All three members of the species complex may be distinguished from each other by 7 − 9 anatomical characters, analysed by MANOVA. Based on 22 enzymes and proteins studied, the moderate to high Wright’s fixation index and the significant (P < 0.05) allele differentiation between EOD phenotypes provide additional evidence for incipient speciation. Pairwise analyses of the three different phenotypes showed the two parapatric species of H. ansorgii grouped together, and distinguishable from individuals of H. szaboi . Analyses of the mitochondrial cytochrome b gene revealed that all specimens which were attributed to H. szaboi form a well-supported monophyletic basal clade (bootstrap support 73% or 82%). The genetic distances (uncorrected p distances) between H. szaboi and the two species of H. ansorgii are between 0.6% and 1.7%. Within the derived H. ansorgii clade some phylogeographical differentiation can be seen for fishes from the Zambezi and Kwando Rivers, but the respective groups are not consistent or supported by significant bootstrap values. The question of which of the two parapatric morphological and EOD phenotypes of H. ansorgii recognized in the present paper represents H. ansorgii (Boulenger, 1905) cannot be resolved at present because of the paucity and unclear origin of the historical type material

What does it feel like to be an electroreceptive fish?

The weakly electric knifefish Eigenmannia emits an electric organ discharge (EOD) of constant frequency and sinusoidal waveform that varies with sex and age. Eigenmannia discriminates among these except when stimulated at the same frequency as its own EOD frequency. In that case, it needs to perform a Jamming Avoidance Response (frequency shift) which results in a beating mixed signal. By a sophisticated analysis of the amplitude and phase modulations of the beat signal, Eigenmannia derives the frequency difference, its sign, and the waveform of the stimulus, hence the signaller’s identity. The human ear is not capable of an equivalent waveform analysis of acoustic stimuli

Allopatric differentiation in the Marcusenius macrolepidotus species complex in southern and eastern Africa: the resurrection of M. pongolensis and M. angolensis, and the description of two new species (Mormyridae, Teleostei)

We critically compared local populations of the bulldog fish, Marcusenius macrolepidotus (Peters 1852), from different watersheds, from the furthest south (28° South, South Africa) to the Equator in Kenya. We ascertained allopatric differentiation from topotypical M. macrolepidotus from the Lower Zambezi River (Mozambique) in morphology, electric organ discharges, and molecular genetics for: (1) samples from the Okavango and Upper Zambezi Systems (Botswana and Namibia), (2) samples from South Africa's rivers draining into the Indian Ocean, and (3) samples from the East African Tana River (Kenya). Significant genetic distances in the mitochondrial cytochrome b gene and differing ISSR-PCR profiles corroborate differentiation between the four taxa. We resurrect M. pongolensis (Fowler, 1934) for South Africa (sample 2), and M. angolensis (Boulenger, 1905) for the Quanza River/Angola. We recognize M. altisambesi sp. n. for the Upper Zambezi/Okavango specimens (sample 1), and M. devosi sp. n. for those from Kenya (sample 3)