Reproduction and development of the weakly electric fish,Pollimyrus isidori (Mormyridae,Teleostei) in captivity

Synopsis No clear sexual dimorphism occurs in Pollimyrus isidori. Females usually grew slightly larger than males. The anal-fin reflex, however, makes it easy to discriminate between males and females. Spawning took place during the first six hours of the dark phase, in the territory of the male. During each spawning act 2–4 eggs were laid. The male put the eggs into a well-hidden nest previously constructed from plant material. The eggs, free embryos, and larvae of several spawnings were guarded by the male for several weeks. The females laid on average 120 eggs per fractional spawning. The eggs were 2 mm in diameter, not adhesive and very yolky. There was a relationship between pH and viability (% embryos hatched). Hatching occurred on the third day after spawning. The respiratory network of segmental origin in the median fin fold was well developed in the free embryos. Transition to exogenous feeding occurred on day 14. The larval period ended when the fish were 15 mm long and 40–50 days old. First gonadal recrudescence occurred at a total length of 6 cm (about 200 days old). The environmental factors decreasing conductivity and pH, increasing water level, and imitation of rain led to gonadal recrudescence, but pH did not act as a cue. Recrudescence was triggered by a decrease of conductivity, but absolute values or ionic composition of the water were not important. Maturation was completed after about fifty days. Mature and spawning fish no longer required any variation of these environmental factors to maintain mature gonads. A steady and considerable increase in conductivity led to gonadal regression. The fish did not show postbreeding refractoriness.     

Acoustic communication in an electric fish,Pollimyrus isidori (Mormyridae)

It has been known since von Frisch's work in the 1930's that mormyrid electric fishes are quite sensitive to sound. We now describe a repertoire of natural sounds produced by the mormyrid, Pollimyrus isidori, during breeding and aggression; reception of communication sounds is probably a major function for mormyrid audition. In aquaria, Pollimyrus isidori produce 'grunts', 'moans', 'growls', 'pops' and 'hoots' at various phases during nesting, courtship, and territory defense. All five sounds are produced primarily at night. Territorial males produce grunts, moans and growls during courtship. Vocalizing is stimulated by the presence of a gravid female on the male's territory and decreases with the onset of spawning. Hoots and pops are given during agonistic behavior. Grunts are bursts of acoustic pulses, stereotyped for an individual, with the potential as individual signatures. The electric organ is silent during grunts and moans and is discharged at a reduced rate during growls. The courtship and spawning of Pollimyrus isidori is described.     

On the electromotor neurons of both electric organs of Pollimyrus isidori (Mormyridae, Teleostei)

The electromotor neurons (EMNs) of the adult electric organ of Pollimyrus isidori and the "giant neurons" (GNs) found in the more rostral part of the spinal cord were investigated both with light and electron microscopical methods. The two kinds of neuron are more or less similar in size, about 30 microns, and are found in the more dorsal part of the spinal cord. The EMNs show somatosomatic gap junctions, as do the GNs. In addition, chemical synapses could be found between nerve fibres and both types of neurons. Histochemical investigations show that the EMNs and the GNs are both AChE positive. On the basis of the arguments presented in the discussion, we believe that the "giant neurons" are the electromotor neurons of the larval electric organ of Pollimyrus isidori.     

Sex differences in the waveform of the pulse-type electric fish,Pollimyrus isidori (Mormyridae)

Summary This paper reports the finding of a sexual dimorphism in the electric organ discharge of the weakly electric fishPollimyms isidori. The difference in the head-positive components of the waveform (Fig. 1) was found in 95% of adults. Juvenile fish always possessed female type discharges. Fourier analysis (Fig. 3) revealed significant sex differences in the power spectra of the discharges, indicating that specific phase information is perhaps unnecessary for sexual identification. The results are discussed with reference to our current knowledge of coding of electrocommunication signals in the electrosensory system.Abbreviation EOD electric organ discharge     

Central auditory neurophysiology of a sound-producing fish: the mesencephalon of Pollimyrus isidori (Mormyridae)

This paper describes the auditory neurophysiology of the mesencephalon of P. isidori, a soundproducing mormyrid fish. Mormyrids have a specialized pressure-sensitive auditory periphery, and anatomical studies indicate that acoustic information is relayed to the mesencephalic nucleus MD. Fish were stimulated with tone bursts and clicks, and responses of MD neurons were recorded extracellularly. Auditory neurons had best frequencies (BF) and best sensitivities (BS) that fell within the range of frequencies and levels of the natural communication sounds of these fish. BSs were in the range of 0 to — 35 dB (re. 1.0 dyne/cm²). Many of the neurons were tuned (Q_{10 dB}: 2–6), and had BFs in the range of 100–300 Hz where the animal's sounds have their peak energy. A range of different physiological cell types were encountered, including phasic, sustained, and complex neurons. Some of the sustained neurons showed strong phase-locking to tones. Many neurons exhibited non-monotonic rate-level functions. Frequencies flanking the BF often caused a reduction in spontaneous activity suggesting inhibition. Many neurons showed excellent representation of click-trains, and some showed a temporal representation of inter-click-intervals with errors less than 1 ms.Abbreviations BF best frequency - BS best sensitivity - ELa anterior exterolateral toral nucleus - ELp posterior exterolateral toral nucleus - EOCD electric organ command discharge - FFT fast Fourier transform - HRP horseradish peroxidase - ICI inter-clickinterval - MD mediodorsal toral nucleus (=auditory nucleus) - OR onset response rate - PSTH peri-stimulus-time-histogram - R synchronization coefficient - RA response area - SS steady state response rate     

Adaptive radiation in African weakly electric fish (Teleostei: Mormyridae: Campylomormyrus): a combined molecular and morphological approach

We combined multiple molecular markers and geometric morphometrics to revise the current taxonomy and to build a phylogenetic hypothesis for the African weakly electric fish genus Campylomormyrus. Genetic data (2039 bp DNA sequence of mitochondrial cytochrome b and nuclear S7 genes) on 106 specimens support the existence of at least six species occurring in sympatry. We were able to further confirm these species by microsatellite analysis at 16 unlinked nuclear loci and landmark-based morphometrics. We assigned them to nominal taxa by comparisons to type specimens of all Campylomormyrus species recognized so far. Additionally, we showed that the shape of the elongated trunk-like snout is the major source of morphological differentiation among them. This finding suggests that the radiation of this speciose genus might have been driven by adaptation to different food sources.     

Trained Weakly-electric Fishes Pollimyrus isidori and Gnathonemus petersii (Mormyridae,Teleostei) Discriminate between Waveforms of Electric Pulse Discharges

Fish of the family Mormyridae emit weak, pulse-like electric organ discharges (EODs). The discharge rhythm is variable, but the waveform of the EOD is constant for each fish, with species- and individual characteristics. The ability of Pollimyrus isidori and Gnathonemus petersii (Mormyridae) to discriminate between different EOD waveforms was tested using a differential conditioning procedure. Fish were first trained to respond to a reference signal in swimming to a dish to receive a bloodworm (food reward). The reference signal consisted of a 10-Hz train of the digitally recorded EOD of a conspecific. Second, an alternative signal (10-Hz train of a different EOD, either from another species, or from a conspecific of the other sex) was associated with air bubbles as punishment. The two signals were played at successive trials in random order. On each trial the latency was measured between the onset of the signal and the response. 7 out of the 8 P. isidori tested and both of the two G. petersii tested associated the reference EOD with food. Among these, five P. isidori and two G. petersii responded differentially (p < 0.01) to EODs of different species. P. isidori similarly discriminated between conspecific EODs of different sexes. The quantity of different alternative EODs which could be tested was limited when fish eventually habituated to the punishment. Even when the amplitude of the EODs was randomly changed at each trial, two out of two G. petersii differentiated between EODs of the two species, and three out of three P. isidori tested differentiated between EODs within their own species. Response latencies to the rewarded signal during the basic training and during discrimination (when it had to be distinguished from the S-) were similar. G. petersii showed differential responses for S+ and S- also in the rhythm of discharge exhibited during playback, after five EOD pulses for one fish, and after a single pulse for the other. Mormyrids may therefore distinguish between conspecifics and members of other species, and even between individual conspecifics, by their EOD waveform.     

Adaptive radiation in the Congo River: An ecological speciation scenario for African weakly electric fish (Teleostei; Mormyridae; Campylomormyrus)

The ultimate aim of this study is to better understand the diversification of African weakly electric fish in the Congo River. Based on a robust phylogenetic hypothesis we examined the radiation within the mormyrid genus Campylomormyrus. Morphological traits relevant for the divergence between the identified species were detected. Among them, the variation in the shape of the trunk-like elongated snout suggested the presence of different trophic specializations. Furthermore, the chosen model taxon, the genus Campylomormyrus, exhibits a wide diversity of electric organ discharge (EOD) waveforms. A comparison of EOD waveform types and phylogenetic relationships showed major differences in EOD between closely related species. This indicates that the EOD might function as a reproductive isolation mechanism. In conclusion, we provide a plausible scenario of an adaptive radiation triggered by sexual selection and assortative mating based on EOD characteristics, but caused by a divergent selection on the feeding apparatus. These findings point towards an adaptive radiation of at least some Campylomormyrus species occurring in the Lower Congo River.     

Electrocommunication and Social Behaviour in Marcusenius senegalensis (Mormyridae,Teleostei)

The electric organ discharges (EODs) of Marcusenius senegalensis, a West African freshwater fish, are bipolar pulses of short duration (220 ± SE 13 μs). In males (n = 10; 10.1–13.1 cm standard length — which is around the size of getting mature), the duration of EOD pulses was of significantly greater variance than in females (n = 9; 9.8–12.8 cm standard length). Male EODs also showed a tendency for a longer duration than female EODs. Groups of three as well as of 14 M. senegalensis formed temporary schools in a ‘naturally’ equipped 720-1 tank. While swimming slowly in a loose school during their nocturnal active phase, fish discharged in irregular long-short-long inter-EOD interval patterns. Near neighbours displayed a tendency to discharge in intervals of similar duration (nearest neighbour distance < 1/2 fish length). On removal of a plastic partition that had separated a pair of fish for at least 3 days, mutual threat displays followed by fighting were observed. During threatening, the fish alternated regularly between bursts of a high discharge rate and short discharge breaks; the rate of change was 4/s. The subdominant animal in a group of two was attacked frequently and often ceased discharging when the dominant fish approached. Courtship behaviour involving gonadally mature fish was accompanied by high-discharge-rate displays with intervals of constant duration in both fish, and the reciprocal display of ‘preferred’ EOD latencies in the 12 ms range. The results demonstrate electric communication by distinct inter-discharge interval patterns in the social behaviour of this mormyrid fish.     

Zur Feinstruktur der Elektrozeptorepidermis der Mormyridae (Teleostei,Pisces)

Zusammenfassung Die schwachelektrischen Mormyridae haben eine dreischichtige Epidermis, deren innere Schicht aus nur etwa 0,22 m dicken sechseckigen Zellen von ca. 60 m Durchmesser besteht. Die etwa 2 m dicken, linsenförmigen Kerne von 7,6 m Durchmesser liegen am Zellrand. Die Zellen sind zu Säulen aufgeschichtet. Ihr Rand ist ausgezackt und dort, wo er die Säulengrenze erreicht, auf etwa 0,34 m verdickt. In der Nähe der Säulengrenzen sind die Zellen über Desmosomen mit den Nachbarn in der eigenen und in der angrenzenden Säule verbunden. Diese Epidermisschicht ist auf die Körperpartien beschränkt, in denen auch Elektrorezeptoren ausgebildet sind.Die beiden anderen Epidermisschichten haben den üblichen Aufbau einer Fischepidermis, abgesehen vom Fehlen der Becherzellen.

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Ultrastructure of the electroceptor epidermis of the Mormyridae (Teleostei, Pisces)

Summary The weakly electric fish of the family Mormyridae have a three layered epidermis, with a medium layer consisting of hexagonal cells of only 0.22 m in thickness and about 60 m in diameter. The lens-shaped nuclei are about 2 m thick and 7.6 m in diameter and are situated near the border of the cells. The cells are piled up to hexagonal columns. Their margin is serrate and where it reaches the boundary of the column, it has a thickness of about 0.34 m. Close to the boundaries of the columns, the cells are linked to their neighbours within the column and of the adjoining column by desmosomes. This layer of the epidermis is confined to those regions of the body surface which also contain electroreceptors.The other layers of the epidermis have a structure as usual in fish, except for the lack of goblet cells.

     

Zum Aufbau der epidermalen Sinnesorgane der Mormyridae (Mormyriformes,Teleostei)

Zusammenfassung Die schwachelektrischen Mormyridae haben außer den elektrischen Organen und ihren oft sehr merkwürdigen Körperformen eine Reihe morphologischer Besonderheiten, die mit ihren elektrischen Fähigkeiten zusammenhängen.Die in der spezifisch gebauten Epidermis liegenden Sinnesorgane, die Mormyromasten, wurden an Hand von Flächenpräparaten untersucht. Die drei Typen, A-, B- und Knollenorgane, sind deutlich voneinander verschieden; sie stellen keineswegs nur Entwicklungsstadien eines Organtyps dar. Mit Toluidinblau ergeben sich sehr auffällige, metachromatische Färbungen.Die Gattung Petrocephalus zeichnet sich durch einen besonderen Typ von Knollenorganen aus, die kleinere, aber zahlreichere Sinneszellen enthalten, als es bei den anderen Gattungen der Fall ist. Sie sind außerdem am Kopf in jederseits drei Rosetten angeordnet, die erstmals beschrieben werden. Die Gefäßversorgung der Mormyromasten wird dargestellt.

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Structure of the epidermal sensory organs in Mormyridae (Mormyriformes, teleostei)

Summary The weakly electric Mormyridae, in addition to their electric organs and their sometimes odd body shapes, have a number of morphological peculiarities which are related to their electric properties.The epidermal sense organs — the mormyromasts —, situated in the specifically structured epidermis, have been investigated by means of surface preparations. The structure of the three types of mormyromasts, A, B and tuberous organs (Knollenorgane) is very different. The mormyromasts are not developmental stages of one type of organ. Toluidine blue stains these sense organs metachromatically.The genus Petrocephalus is characterized by a particular type of tuberous organs, containing smaller but more numerous sensory cells, as compared with other genera. In the head, these organs furthermore are aggregated to three rosettes on either side, described here for the first time.The vascularization of the mormyromasts is demonstrated.

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Mit Unterstützung der Deutschen Forschungsgemeinschaft.     

Hidden species diversity in Marcusenius moorii (Teleostei: Mormyridae) from the Congo Basin

New collections from the Yangambi Biosphere Reserve (YBR) and Okapi Wildlife Reserve (OWR) revealed the presence of two groups of specimens similar to, but different from Marcusenius moorii. To study both these groups, an integrated morphological and genetic (mtDNA, cytb) approach was used. This study revealed that one of the two groups is conspecific with Marcusenius lambouri, a junior synonym of M. moorii, which is herein revalidated, with M. moorii longulus as its junior synonym. Marcusenius lambouri differs from M. moorii by a higher number of lateral line scales (44–46 vs. 40–43), a shorter pectoral-fin length (14.6–19.9 vs. 20.3–25.2% standard length; L_S) and a more elongated body due to a usually shallower middle body depth (19.8–26.5 vs. 26.3–35.9% L_S). The other group revealed to be a new species for science, Marcusenius verheyenorum, which can be distinguished from its congeners with eight circumpeduncular scales by the following unique combination of characters: a rounded head with a terminal mouth; a short and deep caudal peduncle (middle caudal-peduncle depth, 44.9–54.6% caudal-peduncle length; L_CP), a deep body (middle body depth, 27.7–34.2% L_S), 38–43 scales on the lateral line, 40–41 vertebrae, 20–21 dorsal-fin rays and 26 anal-fin rays. Some specimens previously attributed to M. moorii were examined and reassigned to M. lambouri or M. verheyenorum. As a result, M. moorii and M. lambouri occur in sympatry in the middle Congo Basin, with the distribution area of M. moorii still further extending into the lower Congo Basin. Instead, the distribution of M. verheyenorum is limited to some right bank tributaries of the upstream part of the middle Congo Basin. Two museum records from the Lilanda River (YBR), collected in the 1950s and previously identified as M. moorii, were re-identified as belonging to the new species, M. verheyenorum. However, the species now seems locally extinct in that region, which reflects the significant anthropogenic effects even within this reserve.     

Marcusenius desertus sp. nov. (Teleostei: Mormyridae), a mormyrid fish from the Namib desert

We critically compared Marcusenius specimens from the mouth of the Cunene River on the Namibia/Angola border, a harsh desert environment on the Atlantic Ocean coast virtually devoid of aerial insects with aquatic larvae which are an important food item, with Marcusenius multisquamatus Kramer & Wink, 2013 from the escarpment region of that same river, in a relatively rich and productive subtropical savannah environment. River mouth specimens were differentiated in morphology and electric organ discharges, as determined by ANOVA/MANOVA comparisons, principal component and discriminant analyses on morphological and electrophysiological characters, and genetics, including sequences of the mitochondrial cytochrome b gene, indicating reproductive isolation. Specimens from the river mouth differed from M. multisquamatus, their closest relatives, by having a shorter snout, a smaller eye diameter, and smaller nares separation. River mouth specimens were also differentiated from other, increasingly less-close relatives, such as M. altisambesi Kramer et al., 2007 from the Okavango River, Botswana, and from M. krameri Maake et al., 2014 from the Limpopo System, South Africa. We therefore designate the new species Marcusenius desertus sp. nov. for the Cunene River mouth population.     

Electrophysiological and molecular genetic evidence for sympatrically occuring cryptic species in African weakly electric fishes (Teleostei: Mormyridae: Campylomormyrus)

For two sympatric species of African weakly electric fish, Campylomormyrus tamandua and Campylomormyrus numenius, we monitored ontogenetic differentiation in electric organ discharge (EOD) and established a molecular phylogeny, based on 2222bp from cytochrome b, the S7 ribosomal protein gene, and four flanking regions of unlinked microsatellite loci. In C. tamandua, there is one common EOD type, regardless of age and sex, whereas in C. numenius we were able to identify three different male adult EOD waveform types, which emerged from a single common EOD observed in juveniles. Two of these EOD types formed well supported clades in our phylogenetic analysis. In an independent line of evidence, we were able to affirm the classification into three groups by microsatellite data. The correct assignment and the high pairwise F(ST) values support our hypothesis that these groups are reproductively isolated. We propose that in C. numenius there are cryptic species, hidden behind similar and, at least as juveniles, identical morphs.     

Electric organ discharge variability of Mormyridae (Teleostei: Osteoglossomorpha) in the Upper Volta system

The electric organ discharges (EODs) of five mormyrid species ( Marcusenius senegalensis , Brevimyrus niger , Petrocephalus bovei , Pollimyrus isidori , Hippopotamyrus pictus ) from different sampling sites from the Upper Volta system in West Africa were investigated. EOD waveforms were recorded at high sampling rates in order to compare signal waveform parameters of the different species from different locations. Except for H. pictus , EODs within a species differed significantly from one another in some parameters and waveform variability at least between some sampling sites. In addition, each species showed a continuous spectrum of waveform variations, all or only parts of which were found at certain localities. Although there was variability and sometimes similarities between species, the EOD waveforms were species specific. Knowing their variation spectrum, they can be used for species determination and are probably used for species recognition by the mormyrids. Similarities or differences in EOD waveform expression within a species were not related to geographical distance. By contrast, we suggest that biotic environmental factors at a given location influence the expression of EOD waveforms. These factors affect absolute measurements such as EOD duration and fast Fourier transformation peak frequency as well as the amount of variation for certain waveform parameters across species in a similar manner for a given site. Although EOD waveform might be important for the establishment of reproductive barriers between species, our results suggest that differences in waveforms may not necessarily reflect different species or speciation processes in progress.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 61–80.