Mormyrid Species Research Articles

Trophic ecology of the African riverine elephant fishes (Mormyridae).

Multiple species of the elephant fishes (Mormyridae) commonly coexist in sympatry in most African tropical rivers and lakes. In this study, we investigated the trophic ecology and potential trophic niche partitioning of eleven mormyrid fish species from the Sanaga River system in Cameroon using the stable isotope composition of carbon and nitrogen in the muscle samples. Albeit most mormyrids mainly feed on invertebrates, we found differences in isotope ratios, and we report signs of the trophic niche partitioning among species. We further found significant differences in isotopic signatures within the Mormyrus genus, suggesting ecological niche diversification among three closely related species. We have also evaluated differences in the isotopic signals between seasons in four species, which could be possibly caused by species migration and/or anthropogenic agricultural activities. To evaluate body shape, we applied geometric morphometric analyses, and we show that most of the species are clearly morphologically separated. We focused on the mormyrid ecomorphology to identify a possible interaction between shape and ecology, and we found a relationship between the δ13C (but not δ15N) isotopic signal and morphology, suggesting their interplay during mormyrid evolution. Overall, we present robust evidence of the trophic niche partitioning within the mormyrid species community, and we integrate trophic ecology with morphometrics, shedding light on the enigmatic evolutionary history of these fascinating African fishes.

Spike timing-dependent plasticity alters electrosensory neuron synaptic strength in vitro but does not consistently predict changes in sensory tuning in vivo.

How do sensory systems optimize detection of behaviorally relevant stimuli when the sensory environment is constantly changing? We addressed the role of spike timing-dependent plasticity (STDP) in driving changes in synaptic strength in a sensory pathway and whether those changes in synaptic strength could alter sensory tuning. It is challenging to precisely control temporal patterns of synaptic activity in vivo and replicate those patterns in vitro in behaviorally relevant ways. This makes it difficult to make connections between STDP-induced changes in synaptic physiology and plasticity in sensory systems. Using the mormyrid species Brevimyrus niger and Brienomyrus brachyistius, which produce electric organ discharges for electrolocation and communication, we can precisely control the timing of synaptic input in vivo and replicate these same temporal patterns of synaptic input in vitro. In central electrosensory neurons in the electric communication pathway, using whole cell intracellular recordings in vitro, we paired presynaptic input with postsynaptic spiking at different delays. Using whole cell intracellular recordings in awake, behaving fish, we paired sensory stimulation with postsynaptic spiking using the same delays. We found that Hebbian STDP predictably alters sensory tuning in vitro and is mediated by NMDA receptors. However, the change in synaptic responses induced by sensory stimulation in vivo did not adhere to the direction predicted by the STDP observed in vitro. Further analysis suggests that this difference is influenced by polysynaptic activity, including inhibitory interneurons. Our findings suggest that STDP rules operating at identified synapses may not drive predictable changes in sensory responses at the circuit level.NEW & NOTEWORTHY We replicated behaviorally relevant temporal patterns of synaptic activity in vitro and used the same patterns during sensory stimulation in vivo. There was a Hebbian spike timing-dependent plasticity (STDP) pattern in vitro, but sensory responses in vivo did not shift according to STDP predictions. Analysis suggests that this disparity is influenced by differences in polysynaptic activity, including inhibitory interneurons. These results suggest that STDP rules at synapses in vitro do not necessarily apply to circuits in vivo.

Convergence is Only Skin Deep: Craniofacial Evolution in Electric Fishes from South America and Africa (Apteronotidae and Mormyridae).

SynopsisApteronotidae and Mormyridae are species-rich clades of weakly electric fishes from Neotropical and Afrotropical freshwaters, respectively, known for their high morphological disparity and often regarded as a classic example of convergent evolution. Here, we use CT-imaging and 3D geometric morphometrics to quantify disparity in craniofacial morphologies, and to test the hypothesis of convergent skull-shape evolution in a phylogenetic context. For this study, we examined 391 specimens representing 78 species of Apteronotidae and Mormyridae including 30 of 37 (81%) of all valid genera with the goal to sample most of the craniofacial disparity known in these clades. We found no overlap between Apteronotidae and Mormyridae in the skull-shape morphospace using PCA and a common landmark scheme, and therefore no instances of complete phenotypic convergence. Instead, we found multiple potential instances of incomplete convergence, and at least one parallel shift among electric fish clades. The greatest components of shape variance in both families are the same as observed for most vertebrate clades: heterocephaly (i.e., opposite changes in relative sizes of the snout and braincase regions of the skull), and heterorhynchy (i.e., dorsoventral changes in relative snout flexion and mouth position). Mormyrid species examined here exhibit less craniofacial disparity than do apteronotids, potentially due to constraints associated with a larger brain size, ecological constraints related to food-type availability. Patterns of craniofacial evolution in these two clades depict a complex story of phenotypic divergence and convergence in which certain superficial similarities of external morphology obscure deeper osteological and presumably developmental differences of skull form and function. Among apteronotid and mormyrid electric fishes, craniofacial convergence is only skin deep.

Signal Diversification Is Associated with Corollary Discharge Evolution in Weakly Electric Fish.

Communication signal diversification is a driving force in the evolution of sensory and motor systems. However, little is known about the evolution of sensorimotor integration. Mormyrid fishes generate stereotyped electric pulses (electric organ discharge [EOD]) for communication and active sensing. The EOD has diversified extensively, especially in duration, which varies across species from 0.1 to >10 ms. In the electrosensory hindbrain, a corollary discharge that signals the timing of EOD production provides brief, precisely timed inhibition that effectively blocks responses to self-generated EODs. However, corollary discharge inhibition has only been studied in a few species, all with short-duration EODs. Here, we asked how corollary discharge inhibition has coevolved with the diversification of EOD duration. We addressed this question by comparing 7 mormyrid species (both sexes) having varied EOD duration. For each individual fish, we measured EOD duration and then measured corollary discharge inhibition by recording evoked potentials from midbrain electrosensory nuclei. We found that delays in corollary discharge inhibition onset were strongly correlated with EOD duration as well as delay to the first peak of the EOD. In addition, we showed that electrosensory receptors respond to self-generated EODs with spikes occurring in a narrow time window immediately following the first peak of the EOD. Direct comparison of time courses between the EOD and corollary discharge inhibition revealed that the inhibition overlaps the first peak of the EOD. Our results suggest that internal delays have shifted the timing of corollary discharge inhibition to optimally block responses to self-generated signals.SIGNIFICANCE STATEMENT Corollary discharges are internal copies of motor commands that are essential for brain function. For example, corollary discharge allows an animal to distinguish self-generated from external stimuli. Despite widespread diversity in behavior and its motor control, we know little about the evolution of corollary discharges. Mormyrid fishes generate stereotyped electric pulses used for communication and active sensing. In the electrosensory pathway that processes communication signals, a corollary discharge inhibits sensory responses to self-generated signals. We found that fish with long-duration pulses have delayed corollary discharge inhibition, and that this time-shifted corollary discharge optimally blocks electrosensory responses to the fish's own signal. Our study provides the first evidence for evolutionary change in sensorimotor integration related to diversification of communication signals.

Silencing the Spark: CRISPR/Cas9 Genome Editing in Weakly Electric Fish.

Electroreception and electrogenesis have changed in the evolutionary history of vertebrates. There is a striking degree of convergence in these independently derived phenotypes, which share a common genetic architecture. This is perhaps best exemplified by the numerous convergent features of gymnotiforms and mormyrids, two species-rich teleost clades that produce and detect weak electric fields and are called weakly electric fish. In the 50 years since the discovery that weakly electric fish use electricity to sense their surroundings and communicate, a growing community of scientists has gained tremendous insights into evolution of development, systems and circuits neuroscience, cellular physiology, ecology, evolutionary biology, and behavior. More recently, there has been a proliferation of genomic resources for electric fish. Use of these resources has already facilitated important insights with regards to the connection between genotype and phenotype in these species. A major obstacle to integrating genomics data with phenotypic data of weakly electric fish is a present lack of functional genomics tools. We report here a full protocol for performing CRISPR/Cas9 mutagenesis that utilizes endogenous DNA repair mechanisms in weakly electric fish. We demonstrate that this protocol is equally effective in both the mormyrid species Brienomyrus brachyistius and the gymnotiform Brachyhypopomus gauderio by using CRISPR/Cas9 to target indels and point mutations in the first exon of the sodium channel gene scn4aa. Using this protocol, embryos from both species were obtained and genotyped to confirm that the predicted mutations in the first exon of the sodium channel scn4aa were present. The knock-out success phenotype was confirmed with recordings showing reduced electric organ discharge amplitudes when compared to uninjected size-matched controls.

Intraspecific Energetic Trade-Offs and Costs of Encephalization Vary from Interspecific Relationships in Three Species of Mormyrid Electric Fishes

The evolution of increased encephalization comes with an energetic cost. Across species, this cost may be paid for by an increase in metabolic rate or by energetic trade-offs between the brain and other energy-expensive tissues. However, it remains unclear whether these solutions to deal with the energetic requirements of an enlarged brain are related to direct physiological constraints or other evolved co-adaptations. We studied the highly encephalized mormyrid fishes, which have extensive species diversity in relative brain size. We previously found a correlation between resting metabolic rate and relative brain size across species; however, it is unknown how this interspecific relationship evolved. To address this issue, we measured intraspecific variation in relative brain size, the sizes of other organs, metabolic rate, and hypoxia tolerance to determine if intraspecific relationships between brain size and organismal energetics are similar to interspecific relationships. We found that 3 species of mormyrids with varying degrees of encephalization had no intraspecific relationships between relative brain size and relative metabolic rate or relative sizes of other organs, and only 1 species had a relationship between relative brain size and hypoxia tolerance. These species-specific differences suggest that the interspecific relationship between metabolic rate and relative brain size is not the result of direct physiological constraints or strong stabilizing selection, but is instead due to other species level co-adaptations. We conclude that variation within species must be considered when determining the energetic costs and trade-offs underlying the evolution of extreme encephalization.

Phase locking and phase avoidance behavior in eleven Nilotic mormyrid species (Mormyridae, Mormyriformes) in response to external electric stimuli

Latency-dependent responses to artificial electric stimuli in 11 species of the Nilotic mormyrids were studied. In total, more than 350 poststimulus time histograms from 132 individuals were analyzed. All species studied exhibited the latency-dependent responses. Both main types of responses – phase locking and phase avoidance behavior or the preferred latency response (PLR) and preferred latency avoidance (PLA) – were found in eight species (Brevimyrus niger, Hippopotamyrus pictus, Marcusenius cyprinoides, Mormyrus kannume, M. hasselquistii, Petrocephalus bane, P. bovei and ‘Pollimyrus’ petherici), while only PLR were recorded from the remaining three species (Hyperopisus bebe, Mormyrops anguilloides and Mormyrus caschive). In eight species both types of behavior were found in fish of the same sex. In four species both types of behavior were found both in females and males. Finally, in four species both types of behavior were revealed in the same individual (in total, there were eight such individuals). In some individuals of different species the atypical responses with substantially increased latencies (compared to the typical PLR and PLA) were observed. The possible role of the phase locking and phase avoidance behavior in the mormyrid electrolocation and communication is discussed.

The costs of a big brain: extreme encephalization results in higher energetic demand and reduced hypoxia tolerance in weakly electric African fishes.

A large brain can offer several cognitive advantages. However, brain tissue has an especially high metabolic rate. Thus, evolving an enlarged brain requires either a decrease in other energetic requirements, or an increase in overall energy consumption. Previous studies have found conflicting evidence for these hypotheses, leaving the metabolic costs and constraints in the evolution of increased encephalization unclear. Mormyrid electric fishes have extreme encephalization comparable to that of primates. Here, we show that brain size varies widely among mormyrid species, and that there is little evidence for a trade-off with organ size, but instead a correlation between brain size and resting oxygen consumption rate. Additionally, we show that increased brain size correlates with decreased hypoxia tolerance. Our data thus provide a non-mammalian example of extreme encephalization that is accommodated by an increase in overall energy consumption. Previous studies have found energetic trade-offs with variation in brain size in taxa that have not experienced extreme encephalization comparable with that of primates and mormyrids. Therefore, we suggest that energetic trade-offs can only explain the evolution of moderate increases in brain size, and that the energetic requirements of extreme encephalization may necessitate increased overall energy investment.

Cross-tissue and cross-species analysis of gene expression in skeletal muscle and electric organ of African weakly-electric fish (Teleostei; Mormyridae)

BackgroundAfrican weakly-electric fishes of the family Mormyridae are able to produce and perceive weak electric signals (typically less than one volt in amplitude) owing to the presence of a specialized, muscle-derived electric organ (EO) in their tail region. Such electric signals, also known as Electric Organ Discharges (EODs), are used for objects/prey localization, for the identification of conspecifics, and in social and reproductive behaviour. This feature might have promoted the adaptive radiation of this family by acting as an effective pre-zygotic isolation mechanism. Despite the physiological and evolutionary importance of this trait, the investigation of the genetic basis of its function and modification has so far remained limited. In this study, we aim at: i) identifying constitutive differences in terms of gene expression between electric organ and skeletal muscle (SM) in two mormyrid species of the genus Campylomormyrus: C. compressirostris and C. tshokwe, and ii) exploring cross-specific patterns of gene expression within the two tissues among C. compressirostris, C. tshokwe, and the outgroup species Gnathonemus petersii.ResultsTwelve paired-end (100 bp) strand-specific RNA-seq Illumina libraries were sequenced, producing circa 330 M quality-filtered short read pairs. The obtained reads were assembled de novo into four reference transcriptomes. In silico cross-tissue DE-analysis allowed us to identify 271 shared differentially expressed genes between EO and SM in C. compressirostris and C.tshokwe. Many of these genes correspond to myogenic factors, ion channels and pumps, and genes involved in several metabolic pathways. Cross-species analysis has revealed that the electric organ transcriptome is more variable in terms of gene expression levels across species than the skeletal muscle transcriptome.ConclusionsThe data obtained indicate that: i) the loss of contractile activity and the decoupling of the excitation-contraction processes are reflected by the down-regulation of the corresponding genes in the electric organ’s transcriptome; ii) the metabolic activity of the EO might be specialized towards the production and turn-over of membrane structures; iii) several ion channels are highly expressed in the EO in order to increase excitability; iv) several myogenic factors might be down-regulated by transcription repressors in the EO.Electronic supplementary materialThe online version of this article (doi:10.1186/s12864-015-1858-9) contains supplementary material, which is available to authorized users.

Differentiation of morphology, genetics and electric signals in a region of sympatry between sister species of African electric fish (Mormyridae)

Mormyrid fishes produce and sense weak electric organ discharges (EODs) for object detection and communication, and they have been increasingly recognized as useful model organisms for studying signal evolution and speciation. EOD waveform variation can provide important clues to sympatric species boundaries between otherwise similar or morphologically cryptic forms. Endemic to the watersheds of Gabon (Central Africa), Ivindomyrus marchei and Ivindomyrus opdenboschi are morphologically similar to one another. Using morphometric, electrophysiological and molecular characters [cytochrome b sequences and amplified fragment length polymorphism (AFLP) genotypes], we investigated to what extent these nominal mormyrid species have diverged into biological species. Our sampling covered the known distribution of each species with a focus on the Ivindo River, where the two taxa co-occur. An overall pattern of congruence among datasets suggests that I. opdenboschi and I. marchei are mostly distinct. Electric signal analysis showed that EODs of I. opdenboschi tend to have a smaller initial head-positive peak than those of I. marchei, and they often possess a small third waveform peak that is typically absent in EODs of I. marchei. Analysis of sympatric I. opdenboschi and I. marchei populations revealed slight, but significant, genetic partitioning between populations based on AFLP data (F(ST) approximately 0.04). Taken separately, however, none of the characters we evaluated allowed us to discriminate two completely distinct or monophyletic groups. Lack of robust separation on the basis of any single character set may be a consequence of incomplete lineage sorting due to recent ancestry and/or introgressive hybridization. Incongruence between genetic datasets in one individual, which exhibited a mitochondrial haplotype characteristic of I. marchei but nevertheless fell within a genetic cluster of I. opdenboschi based on AFLP genotypes, suggests that a low level of recent hybridization may also be contributing to patterns of character variation in sympatry. Nevertheless, despite less than perfect separability based on any one dataset and inconclusive evidence for complete reproductive isolation between them in the Ivindo River, we find sufficient evidence to support the existence of two distinctive species, I. opdenboschi and I. marchei, even if not 'biological species' in the Mayrian sense.

Sex Discrimination Among Four Mormyrid Species Of Anambra River System Nigeria

Sex discriminating characters of four mormyrid species caught from Anambra river basin, Nigeria were investigated. Sexual dimorphism occurred in only one transformed character – dorsal fin base length and in four raw morphometric characters namely totallength, standard length, dorsal fin base length and anal fin base length. These characters are recommended as key characters in mormyrid taxonomy. Keywords: Mormyridae, sex dimorphism, Anambra river Animal Research International Vol. 1 (3) 2004 pp. 169-172

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.

Length-Weight Relationship And Condition Factor Of Four Mormyrid Species Of Anambra River

The length-weight relatonship and condition factor of four mormyrid species namely Mormyrus rume, Hyperopisus bebe, Campylomormyrus tamandua and Gnathonemus petersii from Anambra river were investigated from October 2002 to March 2004. In al400 species M. rume, 384 H. bebe417 C. tamandua and 335 G. petersii were sampled for the study Length-weigh relationship showed that the exponent “b,\' were 3.067, 2.459, 3.201 and 3.114 for M. rume, H. bebe, C. tamandua and G. petersii respectively. The mormyrid species studied with exception of H. bebe exhibited isometric growth and the correlation coefficients were positive and highly sgnificant (P Keywords : Mormyrids, Length-weight relationship, Condition factor Animal Research International Vol. 3 (1) 2006 pp. 426-430

SOUND PRODUCTION IN THE TERRITORIAL BEHAVIOUR OF THE CHURCHILL PETROCEPHALUS CATOSTOMA (MORMYRIDAE, TELEOSTEI) FROM THE UPPER ZAMBEZI RIVER

ABSTRACT This is the first description of vocalisations produced by the mormyrid species Petrocephalus catostoma from the Upper Zambezi River whilst defending a territory. Agonistic behavioural displays of a dominant male towards a conspecific, such as mutual circling or short attacks, were accompanied by characteristic tonal sounds, termed hoots. The mean hoot duration (43 ± SD 1.8 ms) was longer, and the fundamental frequency (Hi, 180 ± SD 4.7 Hz) lower, than in the closely related species Petrocephalus ballayi. P. catostoma vocalised hoots only during intraspecific agonistic interactions, especially those accompanying territorial conflict.

Larval electroreceptors in the epidermis of mormyrid fish: II. The promormyromast

Promormyromasts were found in the epidermis of the head of the larvae of five species of mormyrids bred in captivity. The promormyromast is a larval electroreceptor belonging to the specific lateral line system. In 12-day-old larvae this electroreceptor is characterized by a single sensory cell and two types of accessory cells. One type of accessory cell has dark cytoplasm, few microtubules, and contacts the sensory cell directly, whereas a second type has pale cytoplasm, many microtubules, and forms an outer layer not directly in contact with the sensory cell. This second type is referred to as a long pyriform accessory cell. This assembly of cells is situated below an intraepidermal cavity filled with acid polysaccharides. The bordering epidermal cells extend microvilli into the intraepidermal cavity. The apexes of the sensory cell, and of the two types of accessory cells, also open into the intraepidermal cavity but bear no microvilli. The promormyromast is innervated by an unmyelinated sensory nerve fiber passing through the basal membrane, which then splits into several branches between the accessory cells. These branches contact the periphery of the sensory cell with terminal boutons. At the site of each contact a ribbon-like structure surrounded by vesicles is present in the cytoplasm of the sensory cell. In older larvae of Campylomormyrus cassaicus, membrane foldings develop at the periphery of the pyriform accessory cells and accessory cell staining properties change just before transformation to become a mormyromast. The functional role of the promormyromast of the larval mormyrids is discussed.

Food habits and trophic resource partitioning among three mormyrid fishes from man-made Lake Ayame, Ivory Coast

ABSTRACT Feeding habits and trophic relationships among three mormyrid species (Petrocephalus bovei, Marcusenius ussheri and Marcusenius furcidens) were assessed in the man-made Lake Ayame in Ivory Coast. The index of preponderance combining the occurrence and weight percentages of the identified prey items showed that these three mormyrid species mainly feed on invertebrates, especially aquatic insects. Petrocephalus bovei mainly feeds on chaoborid larvae, whereas both Marcusenius species preferentially feed on chironomid larvae. This segregation of food items associated with the relative abundance of chironomid larvae may reduce interspecific competition between these three fish species, and may therefore allow their coexistence. Horn's food overlap index revealed highly significant similarities in the trophic spectrum between both Marcusenius species, and for the dry as well as the rainy season.

Food habits and trophic resource partitioning among three mormyrid fishes from man-made Lake Ayame, Ivory Coast

Feeding habits and trophic relationships among three mormyrid species (Petrocephalus bovei, Marcusenius ussheri and Marcusenius furcidens) were assessed in the man-made Lake Ayame in Ivory Coast. The index of preponderance combining the occurrence and weight percentages of the identified prey items showed that these three mormyrid species mainly feed on invertebrates, especially aquatic insects. Petrocephalus bovei mainly feeds on chaoborid larvae, whereas both Marcusenius species preferentially feed on chironomid larvae. This segregation of food items associated with the relative abundance of chironomid larvae may reduce interspecific competition between these three fish species, and may therefore allow their coexistence. Horn’s food overlap index revealed highly significant similarities in the trophic spectrum between both Marcusenius species, and for the dry as well as the rainy season.

Eighteen microsatellite loci for endemic African weakly electric fish (Campylomormyrus, Mormyridae) and their cross species applicability among related taxa

AbstractWe describe isolation and characterization of the first microsatellite loci specifically developed for African weakly electric fish (Mormyridae), for the genus Campylomormyrus. Seventeen of our 18 loci are polymorphic within the Campylomormyrus numenius species complex. The polymorphic loci showed four to 15 alleles per locus, an expected heterozygosity between 0.46 and 0.94, and an observed heterozygosity between 0.31 and 1.00. Most primers also yield reproducible results in several other mormyrid species. These loci comprise a set of molecular markers for various applications, from moderately polymorphic loci suitable for population studies to highly polymorphic loci for pedigree analysis in mormyrids.

SOUND PRODUCTION IN THE REPRODUCTIVE BEHAVIOUR OF THE WEAKLY ELECTRIC FISH POLLIMYR US MARIANNE KRAMER ET AL. 2003 (MORMYRIDAE, TELEOSTEI)

ABSTRACT In addition to their electric organ discharges, two West African mormyrid species are known to generate sounds during reproduction. Here, we document the reproductive behaviour and concurrent vocalisations recorded in captive specimens of the new species Pollimyrus marianne from the Upper Zambezi River. Prior to spawning, three male P. marianne established a territory and built a nest of filamentous plant material each (that would hold eggs and larvae at a later stage). Beginning with territory establishment and usually at night, only the males generated two distinct vocalisations, a tonal moan (average duration, 2.26 ± SD 0.6 s), and a harsh grunt (0.596 ± SD 0.029 s; grunt pulse repetition rate, PRR, 32.1 ± SD 1.24 Hz at 25.4 ± SD 0.15 °C). Both vocalisation types were similar to, but clearly different from, those recorded in the two West African species, P. isidori and P. adspersus, the PRR (Pulse Repetition Rate) of which were higher by 25% and 60%, respectively (at similar temperature). At constant standard length, some of a male P. marianne's sound characteristics varied only slightly over two years, but depended clearly on water temperature. When no other fish were present, the three males produced moans only sporadically. When, however, females were visiting, the males generated long-lasting moan sequences that were superimposed by grunts (moan-grunt bouts; average duration, 14.69 ± SD 2.2 s). The maximum rate of vocalisation occurred in the nights immediately before spawning; this rate receded when larvae were present in the nest, and vocalisations finally stopped after the larvae had dispersed. Vocalisation rate increased again when females restarted to visit the nest site. Long before and also after reproduction, territorial males vocalised with moans and some grunts, when territory intruders evoked escalating agonistic interactions that were correlated with many bites or butts. In agonistic contexts, moan duration, moan peak amplitude frequency and grunt pulse repetition rate differed from the sounds produced during courtship. In P. marianne, the generation of sounds during reproductive and agonistic interactions suggests a dual function: attraction of mates and threatening competing males, or potentially egg-eating females.

Fish Faunal Resurgence in Lake Nabugabo, East Africa

Abstract:In Lake Nabugabo, Uganda, a small satellite of the equatorial Lake Victoria, approximately 50% of the indigenous fish species disappeared from the open waters subsequent to establishment of the introduced predatory Nile perch ( Lates niloticus ). However, several of these species persisted in wetland refugia. Over the past decade, Nile perch in Lake Nabugabo have been intensively fished. Herein we report a resurgence of some indigenous species in open waters. In a multiyear study, we used annual transects in inshore and offshore waters of exposed ( no wetland ) and wetland habitats to document the pattern of resurgence. In 1995, haplochromine cichlids were largely confined to inshore areas, particularly wetland ecotones, and were rare in Nile perch stomachs, as were most other indigenous species. By 2000 haplochromine cichlids were abundant in inshore and offshore areas of both exposed and wetland transects. Several indigenous noncichlids also reappeared in the main lake, including three of the four original mormyrid species. Between 1995 and 1999, there was a dramatic increase in the proportion of haplochromines in the diet of Nile perch. When haplochromines were rare ( 1995 ), Nile perch switched from an invertebrate‐dominated diet to piscivory at a large size ( 30 cm total length ). In 2000, however, Nile perch were strongly piscivorous by 5–10 cm total length. The pattern of faunal loss and recovery in Lake Nabugabo demonstrates the importance of refugia in providing the seeds of resurgence and provides a model with which to understand some changes in Lake Victoria.