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Gymnotiform electric knifefishes are an important yet undersampled component of the Neotropical aquatic biota. We report on the gymnotiform fauna of the Tres Fronteras region located at the triple border of Brazil, Colombia, and Peru in the biodiverse western Amazon. The presence of at least 33 species of gymnotiforms in the Tres Fronteras region is validated from recent sampling efforts and the review of previously collected materials. A key is provided for the identification of the species that have been collected from the region. We comment on the diversity of habitat utilization and intraspecific colour variation of some species. [ABSTRACT FROM AUTHOR]

Abstract The Gymnotiformes, also known as the South American or Neotropical knifefishes, include the strongly electric Electrophorus electricus and many other weakly electric species. These fish possess specialised electric organs that are able to release electric discharges into the water, for electrolocation and communication, and sometimes for predation and defence. All Gymnotiform species possess a myogenic electric organ (mEO) derived from the muscle tissue, and members of the Apteronotidae family uniquely possess a neurogenic electric organ (nEOs) derived from the nervous tissue. A mEO may consist of ‘Type A’ electrocytes that develop within the tail muscle (for example, in Apteronotus leptorhynchus), or ‘Type B’ electrocytes that develop below the tail muscle (for example, in Brachyhypopomus gauderio). In this review, we discuss the diversity in the anatomy, electric discharge and development of electric organs found in different Gymnotiform species, as well as the ecological and environmental factors that have likely contributed to this diversity. We then describe various hypotheses regarding the evolution of electric organs, and discuss the potential evolutionary origin of the nEO: a pair of nerve cords that are located on either side of the aorta in B. gauderio, and which may have expanded and developed into a nEO in the Apteronotidae family during its evolution from a common ancestral species. Finally, we compare potential Gymnotiform phylogenies and their supporting evidence.

The terminal investment hypothesis (TIH) predicts that individuals with favorable prospects for future reproduction (i.e., high residual reproductive value, RRV) should moderate current reproductive investment in favor of growth, survival, and future reproduction, whereas those with low RRV should "terminally invest" by diverting somatic resources towards current reproduction at the expense of future reproduction. However, support for the TIH in wild animal populations is fragmentary, and the ecological contexts of terminal investment remain poorly known. We report a remarkable case of simultaneous terminal investment involving five sympatric species of the electric knifefish genus Brachyhypopomus, from Amazonian floodplain and terra firme stream habitats. We found that terminal investment is synchronized by seasonal breeding, in response to circannual environmental variation in mortality risk. Four species exhibit a uniseasonal iteroparous (annual) life history with complete post‐reproductive mortality after a single breeding season. One species (Brachyhypopomus beebei) exhibits a 2‐year multiseasonal iteroparous life history with breeding in two seasons and post‐reproductive mortality after the second. In mature females and (most) males of the annual species, as well as in both mature female and male second‐year (but not first‐year) B. beebei, we documented an increase in two metrics of reproductive effort (size‐adjusted gonad mass and electric signal amplitude) and a concomitant reduction in somatic condition (size‐adjusted somatic mass), all in response to proximity to the end of the common breeding season, when RRV approximates zero. In mature first‐year B. beebei, we documented neither an increase in reproductive effort nor a decline in somatic condition, implying an alternative strategy of reproductive restraint. Our findings support Kirkwood's disposable soma theory, which posits that death by reproductive exhaustion can be delayed if terminal investment is replaced by reproductive restraint, allowing individuals to survive and breed in a subsequent season. Deferral of the terminal investment response in annual species, and the origin of a gonadal regression‐regeneration sequence, may open pathways for rapid evolutionary transitions to multiseasonal iteroparity. Excepting the age (year‐group) dependency of terminal investment in B. beebei, we were unable to identify intrinsic cues or extrinsic environmental cues for the terminal investment response in Brachyhypopomus. [ABSTRACT FROM AUTHOR]

Brain region size generally scales allometrically with brain size, but mosaic shifts in brain region size independent of brain size have been found in several lineages and may be related to the evolution of behavioral novelty. African weakly electric fishes (Mormyroidea) evolved a mosaically enlarged cerebellum and hindbrain, yet the relationship to their behaviorally novel electrosensory system remains unclear. We addressed this by studying South American weakly electric fishes (Gymnotiformes) and weakly electric catfishes (Synodontis spp.), which evolved varying aspects of electrosensory systems, independent of mormyroids. If the mormyroid mosaic increases are related to evolving an electrosensory system, we should find similar mosaic shifts in gymnotiforms and Synodontis. Using micro-computed tomography scans, we quantified brain region scaling for multiple electrogenic, electroreceptive, and non-electrosensing species. We found mosaic increases in cerebellum in all three electrogenic lineages relative to non-electric lineages and mosaic increases in torus semicircularis and hindbrain associated with the evolution of electrogenesis and electroreceptor type. These results show that evolving novel electrosensory systems is repeatedly and independently associated with changes in the sizes of individual major brain regions independent of brain size, suggesting that selection can impact structural brain composition to favor specific regions involved in novel behaviors.

The Gymnotiformes, also known as the South American or Neotropical knifefishes, include the strongly electric Electrophorus electricus and many other weakly electric species. These fish possess specialised electric organs that are able to release electric discharges into the water, for electrolocation and communication, and sometimes for predation and defence. All Gymnotiform species possess a myogenic electric organ (mEO) derived from the muscle tissue, and members of the Apteronotidae family uniquely possess a neurogenic electric organ (nEOs) derived from the nervous tissue. A mEO may consist of 'Type A' electrocytes that develop within the tail muscle (for example, in Apteronotus leptorhynchus), or 'Type B' electrocytes that develop below the tail muscle (for example, in Brachyhypopomus gauderio). In this review, we discuss the diversity in the anatomy, electric discharge and development of electric organs found in different Gymnotiform species, as well as the ecological and environmental factors that have likely contributed to this diversity. We then describe various hypotheses regarding the evolution of electric organs, and discuss the potential evolutionary origin of the nEO: a pair of nerve cords that are located on either side of the aorta in B. gauderio, and which may have expanded and developed into a nEO in the Apteronotidae family during its evolution from a common ancestral species. Finally, we compare potential Gymnotiform phylogenies and their supporting evidence. [ABSTRACT FROM AUTHOR]

The bluntnose knifefish Brachyhypopomus occidentalis is a primary freshwater fish from north-western South America and Lower Central America. Like other Gymnotiformes, it has an electric organ that generates electric discharges used for both communication and electrolocation. We assembled a high-quality reference genome sequence of B. occidentalis by combining Oxford Nanopore and 10X Genomics linked-reads technologies. We also describe its demographic history in the context of the rise of the Isthmus of Panama. The size of the assembled genome is 540.3 Mb with an N50 scaffold length of 5.4 Mb, which includes 93.8% complete, 0.7% fragmented, and 5.5% of missing vertebrate/Actinoterigie Benchmarking Universal Single-Copy Orthologs. Repetitive elements account for 11.04% of the genome, and 34,347 protein-coding genes were predicted, of which 23,935 have been functionally annotated. Demographic analysis suggests a rapid effective population expansion between 3 and 5 Myr, corresponding to the final closure of the Isthmus of Panama (2.8–3.5 Myr). This event was followed by a sudden and constant population decline during the last 1 Myr, likely associated with strong shifts in both precipitation and sea level during the Pleistocene glacial-interglacial cycles. The de novo genome assembly of B. occidentalis will provide novel insights into the molecular basis of both electric signal productions and detection and will be fundamental for understanding the processes that have shaped the diversity of Neotropical freshwater environments. [ABSTRACT FROM AUTHOR]

Weakly electric gymnotiform fishes use self-generated electric organ discharges (EODs) to navigate and communicate. The electrosensory range for these processes is a function of EOD amplitude, determined by the fish's electric organ (EO) output and the electrical conductivity of the surrounding water. Anthropogenic activity, such as deforestation, dams and industrial/agricultural runoff, are known to increase water conductivity in neotropical habitats, likely reducing the electrosensory range of these fish. We investigated whether fish modulate EO output as means of re-expanding electrosensory range after a rapid increase in water conductivity in the pulse-type Brachyhypopomus gauderio and the wave-type Eigenmannia virescens. Furthermore, because EOD production incurs significant metabolic costs, we assessed whether such compensation is associated with an increase in metabolic rate. Following the conductivity increase, B. gauderio increased EOD amplitude by 20.2±4.3% over 6 days but with no associated increase in metabolic rate, whereas the EOD amplitude of E. virescens remained constant, accompanied by an unexpected decrease in metabolic rate. Our results suggest that B. gauderio uses a compensation mechanism that requires no metabolic investment, such as impedance matching, or a physiological trade-off wherein energy is diverted from other physiological processes to increase EO output. These divergent responses between species could be the result of differences in reproductive life history or evolutionary adaptations to different aquatic habitats. Continued investigation of electrosensory responses to changing water conditions will be essential for understanding the effects of anthropogenic disturbances on gymnotiforms, and potential physiological mechanisms for adapting to a rapidly changing aquatic environment., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2024. Published by The Company of Biologists Ltd.)

Glass knifefish (Eigenmannia) are a group of weakly electric fishes found throughout the Amazon basin. Their electric organ discharges (EODs) are energetically costly adaptations used in social communication and for localizing conspecifics and other objects including prey at night and in turbid water. Interestingly, a troglobitic population of blind cavefish Eigenmannia vicentespelea survives in complete darkness in a cave system in central Brazil. We examined the effects of troglobitic conditions, which includes a complete loss of visual cues and potentially reduced food sources, by comparing the behavior and movement of freely behaving cavefish to a nearby epigean (surface) population (Eigenmannia trilineata). We found that the strengths of electric discharges in cavefish were greater than in surface fish, which may result from increased reliance on electrosensory perception, larger size, and sufficient food resources. Surface fish were recorded while feeding at night and did not show evidence of territoriality, whereas cavefish appeared to maintain territories. Surprisingly, we routinely found both surface and cavefish with sustained differences in EOD frequencies that were below 10 Hz despite being within close proximity of about 50 cm. A half century of analysis of electrosocial interactions in laboratory tanks suggest that these small differences in EOD frequencies should have triggered the “jamming avoidance response,” a behavior in which fish change their EOD frequencies to increase the difference between individuals. Pairs of fish also showed significant interactions between EOD frequencies and relative movements at large distances, over 1.5 m, and at high differences in frequencies, often >50 Hz. These interactions are likely “envelope” responses in which fish alter their EOD frequency in relation to higher order features, specifically changes in the depth of modulation, of electrosocial signals.

Glass knifefish (Eigenmannia) are a group of weakly electric fishes found throughout the Amazon basin. Their electric organ discharges (EODs) are energetically costly adaptations used in social communication and for localizing conspecifics and other objects including prey at night and in turbid water. Interestingly, a troglobitic population of blind cavefish Eigenmannia vicentespelea survives in complete darkness in a cave system in central Brazil. We examined the effects of troglobitic conditions, which includes a complete loss of visual cues and potentially reduced food sources, by comparing the behavior and movement of freely behaving cavefish to a nearby epigean (surface) population (Eigenmannia trilineata). We found that the strengths of electric discharges in cavefish were greater than in surface fish, which may result from increased reliance on electrosensory perception, larger size, and sufficient food resources. Surface fish were recorded while feeding at night and did not show evidence of territoriality, whereas cavefish appeared to maintain territories. Surprisingly, we routinely found both surface and cavefish with sustained differences in EOD frequencies that were below 10 Hz despite being within close proximity of about 50 cm. A half century of analysis of electrosocial interactions in laboratory tanks suggest that these small differences in EOD frequencies should have triggered the "jamming avoidance response," a behavior in which fish change their EOD frequencies to increase the difference between individuals. Pairs of fish also showed significant interactions between EOD frequencies and relative movements at large distances, over 1.5 m, and at high differences in frequencies, often >50 Hz. These interactions are likely "envelope" responses in which fish alter their EOD frequency in relation to higher order features, specifically changes in the depth of modulation, of electrosocial signals. [ABSTRACT FROM AUTHOR]

Fourteen novel microsatellite loci are described and characterized in two species of electric eels, Electrophorus varii and E. voltai from floodplains and rivers of the Amazon rainforest. These loci are polymorphic, highly informative, and have the capacity to detect reliable levels of genetic diversity. Likewise, the high combined probability of paternity exclusion value and low combined probability of genetic identity value obtained demonstrate that the new set of loci displays suitability for paternity studies on electric eels. In addition, the cross-amplification of electric eel species implies that it may also be useful in the study of the closely related E. electricus, and to other Neotropical electric fishes (Gymnotiformes) species as tested herein. [ABSTRACT FROM AUTHOR]

We quantified patterns of circannual reproductive breeding periodicity in a multi-species assemblage of the gymnotiform knifefish genus Brachyhypopomus from Amazonian floodplain and terra firme stream habitats. We applied linear mixed-effect models and model selection to a dataset of 3205 fish from the five most abundant species to identify environmental variables predictive of gonadosomatic index (GSI) – including rainfall, river level, water temperature, conductivity, dissolved oxygen (DO), and pH. A single common floodplain species exhibited a well-demarcated ca. 6-month breeding season corresponding to the rising-water and high-water period. In this species the strongest predictors of GSI in both sexes were river level and pH. Conductivity, temperature, and DO all exhibited substantial circannual variation in the floodplain but were uncorrelated to GSI. Four common terra firme stream-dwelling species exhibited longer (ca. 7–9-month) and less distinct breeding seasons that were approximately coincident with the late dry/early rainy season. The stream species exhibited a congruent species-wide response in which GSI was predicted in both sexes by rainfall, but not by physico-chemical water properties (all of which exhibited relatively minor circannual variation). Five of the most abundant species in this study have a semelparous one-year life history, but one (B. beebei) has an iteroparous two-year life history. Consistent with a pattern of terminal reproductive investment, we observed a more pronounced elevation of GSI (especially in females) in B. beebei's final second breeding season than in its first. [ABSTRACT FROM AUTHOR]

ABSTRACT We describe the circuit design, construction, and operation of a field-portable electric fish finder (an AC-coupled wide-band differential bio-amplifier with loudspeaker output). This device permits detection and monitoring of the electric organ discharges generated by neotropical gymnotiform fishes (as well as the mormyroid fishes of tropical Africa). Our design is modified from earlier versions to optimize detection performance and stability over a wider range of ambient water conductivity, including under conditions of extremely low conductivity (< ca. 10 μScm-1). Our new electric fish finder design also incorporates complete waterproofing and longer battery autonomy. We provide Gerber and Eagle files made with the electronic design automation software ‘Autodesk Eagle’ to allow researchers to order printed circuit boards directly from commercial manufacturers.

Eavesdropping by electroreceptive predators poses a conflict for weakly electric fish, which depend on their Electric Organ Discharge (EOD) signals both for navigation and communication in the dark. The EODs that allow weakly electric fish to electrolocate and communicate in the dark may attract electroreceptive predators such as catfishes and Electric Eels. These predators share with their prey the synapomorphy of passive electric sense supported by ampullary electroreceptors that are highly sensitive to low-frequency electric fields. Any low-frequency spectral components of the EOD make weakly electric fish conspicuous and vulnerable to attack from electroreceptive predators. Accordingly, most weakly electric fish shift spectral energy upwards or cloak low-frequency energy with compensatory masking signals. Subadults and females in particular emit virtually no low-frequency energy in their EODs, whereas courting males include a significant low-frequency component, which likely attracts females, but makes the signals conspicuous to predators. Males of species that coexist with the most predators tend to produce the least low-frequency signal energy, expressing sexual dimorphism in their signals in less risky ways. In these respects, electric signals follow the classic responses to opposing forces of natural and sexual selection, as exemplified in the visual signals of guppies and the acoustic signals of Túngara frogs. Unique to electric fish is that the electric signal modifications that help elude detection by electroreceptive predators are additions to the basal signal rather than losses of attractive components. These enhancements that enable crypsis are energetically costly, but have also provided the evolutionary substrates for subsequent sexual selection and species identity characters.

Hypoxia is a significant source of metabolic stress that activates many cellular pathways involved in cellular differentiation, proliferation, and cell death. Hypoxia is also a major component in many human diseases and a known driver of many cancers. Despite the challenges posed by hypoxia, there are animals that display impressive capacity to withstand lethal levels of hypoxia for prolonged periods of time and thus offer a gateway to a more comprehensive understanding of the hypoxic response in vertebrates. The weakly electric fish genus Brachyhypopomus inhabits some of the most challenging aquatic ecosystems in the world, with some species experiencing seasonal anoxia, thus providing a unique system to study the cellular and molecular mechanisms of hypoxia tolerance. In this study, we use closely related species of Brachyhypopomus that display a range of hypoxia tolerances to probe for the underlying molecular mechanisms via hypoxia inducible factors (HIFs)-transcription factors known to coordinate the cellular response to hypoxia in vertebrates. We find that HIF1⍺ from hypoxia tolerant Brachyhypopomus species displays higher transactivation in response to hypoxia than that of intolerant species, when overexpressed in live cells. Moreover, we identified two SUMO-interacting motifs near the oxygen-dependent degradation and transactivation domains of the HIF1⍺ protein that appear to boost transactivation of HIF1, regardless of the genetic background. Together with computational analyses of selection, this shows that evolution of HIF1⍺ are likely to underlie adaptations to hypoxia tolerance in Brachyhypopomus electric fishes, with changes in two SUMO-interacting motifs facilitating the mechanism of this tolerance., Competing Interests: Conflict of interest The authors declare that they have no conflicts of interest with the contents of this article., (Copyright © 2024 The Authors. Published by Elsevier Inc. All rights reserved.)

The South American weakly electric fish, Gymnotus omarorum, displays territorial aggression year-round in both sexes. To examine the role of rapid androgen modulation in non-breeding aggression, we administered acetate cyproterone (CPA), a potent inhibitor of androgen receptors, to both male and females, just before staged agonistic interactions. Wild-caught fish were injected with CPA and, 30 min later, paired in intrasexual dyads. We then recorded the agonistic behavior which encompasses both locomotor displays and emission of social electric signals. We found that CPA had no discernible impact on the levels of aggression or the motivation to engage in aggressive behavior for either sex. However, CPA specifically decreased the expression of social electric signals in both males and female dyads. The effect was status-dependent as it only affected subordinate electrocommunication behavior, the emission of brief interruptions in their electric signaling ("offs"). This study is the first demonstration of a direct and rapid androgen effect mediated via androgen receptors on non-breeding aggression. Elucidating the mechanisms involved in non-breeding aggression in this teleost model allows us to better understand potentially conserved or convergent neuroendocrine mechanisms underlying aggression in vertebrates., (Copyright © 2023. Published by Elsevier Inc.)

Active electroreception-the ability to detect objects and communicate with conspecifics via the detection and generation of electric organ discharges (EODs)-has evolved convergently in several fish lineages. South American electric fishes (Gymnotiformes) are a highly species-rich group, possibly in part due to evolution of an electric organ (EO) that can produce diverse EODs. Neofunctionalization of a voltage-gated sodium channel gene accompanied the evolution of electrogenic tissue from muscle and resulted in a novel gene (scn4aa) uniquely expressed in the EO. Here, we investigate the link between variation in scn4aa and differences in EOD waveform. We combine gymnotiform scn4aa sequences encoding the C-terminus of the Na v 1.4a protein, with biogeographic data and EOD recordings to test whether physiological transitions among EOD types accompany differential selection pressures on scn4aa. We found positive selection on scn4aa coincided with shifts in EOD types. Species that evolved in the absence of predators, which likely selected for reduced EOD complexity, exhibited increased scn4aa evolutionary rates. We model mutations in the protein that may underlie changes in protein function and discuss our findings in the context of gymnotiform signalling ecology. Together, this work sheds light on the selective forces underpinning major evolutionary transitions in electric signal production.

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Functional variation in rhodopsin, the dim-light-specialized visual pigment, frequently occurs in species inhabiting light-limited environments. Variation in visual function can arise through two processes: relaxation of selection or adaptive evolution improving photon detection in a given environment. Here, we investigate the molecular evolution of rhodopsin in Gymnotiformes, an order of mostly nocturnal South American fishes that evolved sophisticated electrosensory capabilities. Our initial sequencing revealed a mutation associated with visual disease in humans. As these fishes are thought to have poor vision, this would be consistent with a possible sensory trade-off between the visual system and a novel electrosensory system. To investigate this, we surveyed rhodopsin from 147 gymnotiform species, spanning the order, and analysed patterns of molecular evolution. In contrast with our expectation, we detected strong selective constraint in gymnotiform rhodopsin, with rates of non-synonymous to synonymous substitutions lower in gymnotiforms than in other vertebrate lineages. In addition, we found evidence for positive selection on the branch leading to gymnotiforms and on a branch leading to a clade of deep-channel specialized gymnotiform species. We also found evidence that deleterious effects of a human disease-associated substitution are likely to be masked by epistatic substitutions at nearby sites. Our results suggest that rhodopsin remains an important component of the gymnotiform sensory system alongside electrolocation, and that photosensitivity of rhodopsin is well adapted for vision in dim-light environments. [ABSTRACT FROM AUTHOR]

The most broadly expressed and studied aspect of sensory transduction is receptor tuning to the power spectral density of the incoming signals. Temporal cues expressed in the phase spectrum are relevant in African and American pulse-emitting electric fish showing electroreceptors sensing the signals carried by the selfand conspecific-generated electric organ discharges. This article concerns the role of electroreceptor phase sensitivity in American pulse Gymnotiformes. These fish show electroreceptors sharply tuned to narrow frequency bands. This led to the common thought that most electrosensory information is contained in the amplitude spectra of the signals. However, behavioral and modeling studies suggest that in their pulses, Gymnotiformes electroreceptors also encode cues embodied in the phase spectrum of natural stimuli. Here, we show that the two main types of tuberous primary afferents of Gymnotus omarorum differentially respond to cues embodied in the amplitude and phase spectra of self-generated electrosensory signals. One afferent type, pulse markers, is mainly driven by the amplitude spectrum, while the other, burst coders, is predominantly sensitive to the phase spectrum. This dual encoding strategy allows the fish to create a sensory manifold where patterns of 'electric color' generated by object impedance and other potential sources of 'colored' images (such as large nearby objects and other electric fish) can be represented. [ABSTRACT FROM AUTHOR]

Abstract Electric fishes are a diverse group of freshwater organisms with the ability to generate electric organ discharges (EODs) that are used for communication and electrolocation. This group (ca. 200 species) has originated in South America, and six species colonized the Central American Isthmus. Here, we assembled the complete mitochondrial genomes (mitogenomes) for three Central American electric fishes (i.e. Sternopygus dariensis , Brachyhypopomus occidentalis , and Apteronotus rostratus), and, based on these data, explored their phylogenetic position among Gymnotiformes. The three mitogenomes show the same gene order, as reported for other fishes, with a size ranging from 16,631 to 17,093 bp. We uncovered a novel 60 bp intergenic spacer (IGS) located between the COII and tRNALys genes, which appears to be unique to the Apteronotidae. Furthermore, phylogenetic relationships supported the traditional monophyly of Gymnotiformes, with the three species positioned within their respective family. In addition, the genus Apteronotus belongs to the early diverging lineage of the order. Finally, we found high sequence divergence (13%) between our B. occidentalis specimen and a sequence previously reported in GenBank, suggesting that the prior mitogenome of B. occidentalis represents a different South American species. Indeed, phylogenetic analyses using Cytochrome b gene across the genus placed the previously reported individual within B. bennetti. Our study provides novel mitogenome resources that will advance our understanding of the diversity and phylogenetic history of Neotropical fishes. Highlights • The mitochondrial genomes of Apteronotus rostratus , Brachyhypopomus occidentalis and Sternopygus dariensis are sequenced and characterized. • A novel 60 bp intergenic spacer located between COII and tRNALys is reported for the first time in Gymnotiformes, and it may be unique to Apteronotus. • Phylogenetic analyses support the position of Central American weakly-electric fishes within the monophyletic Gymnotiformes. • Genetic analyses indicate that the mitogenome of B. occidentalis (AP011570) reported previously belongs to a different South America species. [ABSTRACT FROM AUTHOR]

Complete mitochondrial genomes of the characiform fishes Astyanax fasciatus, Astyanax altiparanae, Hoplias malabaricus (Karyomorph A) and the Gymnotiformes species Gymnotus sylvius and Gymnotus cuia were characterized in the present study. The whole mitogenomes varied from 16,400bp (A. fasciatus) to 17,730 bp (A. altiparanae) long and all of them consisted of 13 protein-coding genes, 22 tRNAs, 2 rRNAs genes, a control region, and origin of light-strand replication. The gene order was similar among all the analyzed species. The nucleotide content of all mitogenomes was also similar, with 29.58–30.95% for A, 27.02–28.65% for T, 26.29–29.99% for C, and 14.41–15.67% for G.

Few reports are available describing lesions in captive electric eels Electrophorus spp. This report describes 2 types of cutaneous proliferative lesions (i.e. hamartoma and neoplasm) in a captive electric eel. Ampullary electroreceptor hamartomas appeared grossly as 2 discrete, smooth, pink, spherical, cutaneous masses measuring 6 and 18 mm in diameter. Histologically, hamartomas were composed of predominately spindle cells that were separated into lobules by a peripheral rim of polygonal cells. Spindle cells were arranged in vague streams and occasionally whorls within a myxomatous matrix. Polygonal cells arranged in variably sized trabeculae and cords within a pre-existing fibrovascular stroma surrounded the streams of spindle cells. Admixed with the polygonal cell population were multiple mucous glands and alarm cells, similar to those seen in normal regions of epidermis. Histochemical stains confirmed similar components in the normal ampullary electroreceptor as in the hamartomas. Lymphoma was also present, appearing grossly as patchy pitting, erythematous, and thickened areas of the skin affecting the entire animal. Lymphoma was diffusely infiltrating and expanding the epidermis, oral mucosa, and branchial mucosa up to 1.5 mm in thickness. It was composed of an unencapsulated, well-demarcated, moderately cellular neoplasm composed of lymphocytes arranged in small dense sheets and clusters that separated and effaced epidermal cells. This is the first report of lymphoma in an electric eel, and the first report of ampullary electroreceptor hamartoma in any animal species.

The electric organ discharges (EODs) produced by weakly electric fish have long been a source of scientific intrigue and inspiration. The study of these species has contributed to our understanding of the organization of fixed action patterns, as well as enriching general imaging theory by unveiling the dual impact of an agent's actions on the environment and its own sensory system during the imaging process. This Centenary Review firstly compares how weakly electric fish generate species- and sex-specific stereotyped electric fields by considering: (1) peripheral mechanisms, including the geometry, channel repertoire and innervation of the electrogenic units; (2) the organization of the electric organs (EOs); and (3) neural coordination mechanisms. Secondly, the Review discusses the threefold function of the fish-centered electric fields: (1) to generate electric signals that encode the material, geometry and distance of nearby objects, serving as a short-range sensory modality or 'electric touch'; (2) to mark emitter identity and location; and (3) to convey social messages encoded in stereotypical modulations of the electric field that might be considered as species-specific communication symbols. Finally, this Review considers a range of potential research directions that are likely to be productive in the future., Competing Interests: Competing interests The author declares no competing or financial interests., (© 2023. Published by The Company of Biologists Ltd.)

Neotropical, nocturnal freshwater knifefishes of the families Sternopygidae and Apteronotidae are electroreceptive, and emit electric organ discharges (EODs) of the wave type for communication and active electrolocation. In a field-collected sample of an estimated 43 gymnotiform species, members of the former family displayed the same type of sinusoidal EOD waveform at frequencies of up to about 800 Hz, with the fundamental frequency, f1, the strongest harmonic in each. Members of the latter (Apteronotidae) displayed f1 frequencies of up to 1800 Hz, and a great diversity in EOD waveform. Apteronotid EODs often differed from those of sternopygids by more harmonics at stronger amplitudes, where f1 was not always the strongest harmonic. The frequency band at −10 dB increased with diminishing f1 amplitude. In contrast to apteronotids, all sternopygids showed the same phase relationship between their respective f1 and f2: a difference of an average 72°, which explains their single type of sinusoidal waveform. In apteronotids a great variety of phase relationships among harmonics was observed, in some their harmonics series cycled through 360° repeatedly. It is argued that the evolutionary driving force for the apteronotids — in contrast to sternopygids — was the greater potential for adaptive radiation. [ABSTRACT FROM AUTHOR]

Convergent evolution of a novel locomotor strategy implies that a fitness benefit may be associated with the new gait. Opportunities to study this phenomenon are often constrained by a lack of transitional taxa, but teleost fishes offer examples of extant species across such evolutionary shifts in gait. For instance, one species from Osteoglossiformes and the entire order of Gymnotiformes independently evolved a novel gait, gymnotiform locomotion, where thrust is produced by the undulation of an elongate anal fin. Here, we investigate whether this convergence in gait is also associated with similarities in shape, burst swimming abilities, and/or steady‐swimming energetics. Specifically, we measured body and fin morphology of fish within Gymnotiformes and Osteoglossiformes, along with closely related Siluriformes and Cypriniformes, to examine the link between gymnotiform locomotion and morphology in a phylogenetic context. Second, we tested the burst swimming capabilities and oxygen consumption during endurance swimming of a subset of the same gymnotiform, osteoglossiform, and cypriniform species, including "transitional" Osteoglossiformes that exhibit intermediate gaits, to determine whether the evolution of this specialized gait is associated with a change in either of these performance metrics. Our results suggest that convergence on the gymnotiform gait is associated with morphological convergence, but does not constrain a fish's maximum sprinting speeds or their energetic demands during steady swimming. [ABSTRACT FROM AUTHOR]

We describe the anatomy and histology of the accessory electric organs of several knifefish taxa. Accessory electric organs are observed among Rhamphichthyoidea in the opercular, mental and humeral regions. Within this group, some species of Brachyhypopomus possess an accessory electric organ in the opercular region. Rhamphichthyinae and Steatogenys possess accessory electric organs in the mental region of the body that differs in many aspects, such as general electrocyte shape and its number of caudal ridges. Steatogenys, Hypopygus and Rhamphichthys possess an accessory electric organ in the humeral region that differs in position, electrocyte configuration and shape. Electrocytes of both humeral and mental accessory electric organs in Steatogenys share a number of common features (e.g., electrocyte shape and innervation pattern), which distinguishes them from the electric organs of related groups. Rhamphichthys has an accessory electric organ in the humeral (specifically subpectoral) region, which has not previously been reported in the literature and differs in arrangement and electrocyte shape from those previously described electric organs of other taxa. Homology of these accessory electric organs is discussed in the context of hypothesized relationships among rhamphichthyoid taxa, indicating that accessory electric organs originated multiple times with apparently no subsequent losses. [ABSTRACT FROM AUTHOR]

Chromosome changes can perform an important role in speciation by acting as postzygotic reproductive barriers. The Neotropical electric fish genus Brachyhypopomus (Gymnotiformes, Hypopomidae) has 28 described species, but cytogenetic data are hitherto available only for four of them. To understand karyotype evolution and investigate the possible role of chromosome changes in the diversification of this genus, we describe here the karyotype of eight species of Brachyhypopomus from a sympatric assemblage in the central Amazon basin. We analyzed cytogenetic data in the context of a phylogenetic reconstruction of the genus and known patterns of geographical distribution. We found a strong phylogenetic signal for chromosome number and noted that sympatric species have exclusive karyotypes. Additional insights into the role of chromosome changes in the diversification of Brachyhypopomus are discussed. [ABSTRACT FROM AUTHOR]

Abstract The aim of this study was to determine the morphometric measures and morphological aspects of nematode-cysts in Gymnotus inaequilabiatus, and the presence of melanomacrophage centers (MMCs) associated with the periphery of cysts and in the liver parenchyma. Adult specimens, 34 female (123.1 ± 43.9g) and 45 male (135.5 ± 43.4g), from Paraguay River, Corumbá, Brazil, were used. The number of nematode-cysts was determined in 79 livers and 25 of them randomly selected for histopathological analysis and morphometric measures of nematode-cysts (mean diameter, thickness of collagen layer, and cyst-wall layer). The percentage of cysts with MMCs on the periphery and density in the liver parenchyma was estimated. The average number of macroscopic cysts was of 48.7 ± 2.78. Granulomatous reaction was observed surrounding the cysts. Diameter, collagen layer and cyst-wall measurements were 293.0 ± 75.18 (µm), 17.72 ± 6.01 (µm) and 12.21 ± 9.51 (µm), respectively. The number of nematode-cysts was correlated with hepatosomatic index, (r=0.26, P

The genus Gymnotus (Gymnotiformes) contains over 40 species of freshwater electric fishes exhibiting a wide distribution throughout Central and South America, and being particularly prevalent in the Amazon basin. Cytogenetics has been an important tool in the cytotaxonomy and elucidation of evolutionary processes in this genus, including the unraveling the variety of diploid chromosome number (2n = from 34 to 54), the high karyotype diversity among species with a shared diploid number, different sex chromosome systems, and variation in the distribution of several Repetitive DNAs and colocation and association between those sequences. Recently whole chromosome painting (WCP) has been used for tracking the chromosomal evolution of the genus, showing highly reorganized karyotypes and the conserved synteny of the NOR bearing par within the clade G. carapo. In this study, painting probes derived from the chromosomes of G. carapo (GCA, 2n = 42, 30 m/sm + 12 st/a) were hybridized to the mitotic metaphases of G. arapaima (GAR, 2n = 44, 24 m/sm + 20 st/a). Our results uncovered chromosomal rearrangements and a high number of repetitive DNA regions. From the 12 chromosome pairs of G. carapo that can be individually differentiated (GCA1-3, 6, 7, 9, 14, 16, and 18-21), six pairs (GCA 1, 9, 14, 18, 20, 21) show conserved homology with GAR, five pairs (GCA 1, 9, 14, 20, 21) are also shared with cryptic species G. carapo 2n = 40 (34 m/sm + 6 st/a) and only the NOR bearing pair (GCA 20) is shared with G. capanema (GCP 2n = 34, 20 m/sm + 14 st/a). The remaining chromosomes are reorganized in the karyotype of GAR. Despite the close phylogenetic relationships of these species, our chromosome painting studies demonstrate an extensive reorganization of their karyotypes. [ABSTRACT FROM AUTHOR]

This paper presents data collected in the Upper Maroni, French Guiana during 5 field missions from 2000 to 2002. A total of 123 known species, classified into 34 families, was captured. Fish communities sampled vary during the year depending on the season. On a daily cycle, fish composition also clearly varies, with a predominance of Siluriformes and Gymnotiformes during the night. Our results give a historical overview of the fish fauna in Upper Maroni 20 years ago. At this time, threatened fishes were already recorded due to overfishing and in certain areas, habitat alteration (turbidity and pollution). [ABSTRACT FROM AUTHOR]

The deep channels of large rivers throughout the humid Neotropics are occupied by diverse and abundant assemblages of electric knifefishes. Historically this habitat has been poorly sampled, but extensive benthic trawling efforts in the Brazilian Amazon in the 1990s produced large numbers of electric fishes especially in the family Apteronotidae. A large number of these specimens, initially identified as Porotergus, have been found to belong within Compsaraia, a genus with two species described from the Orinoco and western Amazon. From this material we describe a new species, from the Amazon River in Brazil, and provide a new diagnosis for the genus. This species is readily distinguished from congeners by a short, rounded snout and small, subterminal mouth with reduced dentition. This species inhabits large rivers in the Eastern and Central Amazon between Ilha Grande de Gurupá and the mouth of the Rio Içá. This description brings the total number of valid apteronotid species to 95. [ABSTRACT FROM AUTHOR]

Fishes of the order Gymnotiformes have high diversity of oral and head morphology, which suggests trophic specializations within each clade. The aim of this study was to describe resource use patterns by two fish species (Gymnorhamphichthys rondoni and Gymnotus coropinae) in the National Forest Saracá-Taquera, Oriximiná - Pará, analyzing microhabitat use, diet composition, feeding strategies, niche breadth and niche overlap. Stomach contents of 101 individuals (41 G. rondoni and 60 G. coropinae), sampled in 23 headwater streams were analyzed and volume of food items was quantified to characterize their feeding ecology. Gymnorhamphichthys rondoni was captured mainly on sandy bottoms, whereas G. coropinae in crevices. Both species had a zoobenthivorous diet and consumed predominantly Sediment/Detritus and Diptera larvae, but also included allochthonous prey in their diet. These species had high niche overlap, with small variations related to the higher consumption of Ceratopogonidae larvae by G. rondoni and of Chironomidae larvae by G. coropinae. Both species had a generalist feeding strategy, but G. coropinae had a broader niche breadth. Our results demonstrate that G. rondoni and G. coropinae occupy different microhabitats but rely on similar food resources. [ABSTRACT FROM AUTHOR]

Background: Nocturnally active gymnotiform weakly electric fish generate electric signals for communication and navigation, which can be energetically taxing. These fish mainly inhabit the Amazon basin, where some species prefer well-oxygenated waters and others live in oxygen-poor, stagnant habitats. The latter species show morphological, physiological, and behavioral adaptations for hypoxia-tolerance. However, there have been no studies of hypoxia tolerance on the molecular level. Globins are classic respiratory proteins. They function principally in oxygenbinding and -delivery in various tissues and organs. Here, we investigate the molecular evolution of alpha and beta hemoglobins, myoglobin, and neuroglobin in 12 gymnotiforms compared with other teleost fish. Results: The present study identified positively selected sites (PSS) on hemoglobin (Hb) and myoglobin (Mb) genes using different maximum likelihood (ML) methods; some PSS fall in structurally important protein regions. This evidence for the positive selection of globin genes suggests that the adaptive evolution of these genes has helped to enhance the capacity for oxygen storage and transport. Interestingly, a substitution of a Cys at a key site in the obligate air-breathing electric eel (Electrophorus electricus) is predicted to enhance oxygen storage of Mb and contribute to NO delivery during hypoxia. A parallel Cys substitution was also noted in an air-breathing African electric fish (Gymnarchus niloticus). Moreover, the expected pattern under normoxic conditions of high expression of myoglobin in heart and neuroglobin in the brain in two hypoxia-tolerant species suggests that the main effect of selection on these globin genes is on their sequence rather than their basal expression patterns. Conclusion: Results indicate a clear signature of positive selection in the globin genes of most hypoxia-tolerant gymnotiform fishes, which are obligate or facultative air breathers. These findings highlight the critical role of globin genes in hypoxia tolerance evolution of Gymnotiform electric fishes. [ABSTRACT FROM AUTHOR]

An apteronotid weakly electric fish, Adontosternarchus, emits high-frequency electric organ discharges (700-1500 Hz) which are stable in frequency if no other fish or artificial signals are present. When encountered with an artificial signal of higher frequency than the fish's discharge, the fish raised its discharge frequency and eventually matched its own frequency to that of the artificial signal. At this moment, phase locking was observed, where the timing of the fish's discharge was precisely stabilized at a particular phase of the artificial signal over a long period of time (up to minutes) with microsecond precision. Analyses of the phase-locking behaviors revealed that the phase values of the artificial stimulus at which the fish stabilizes the phase of its own discharge (called lock-in phases) have three populations between −180° and +180°. During the frequency rise and the phase-locking behavior, the electrosensory system is exposed to the mixture of feedback signals from its electric organ discharges and the artificial signal. Since the signal mixture modulates in both amplitude and phase, we explored whether amplitude or phase information participated in driving the phase-locking behavior, using a numerical model. The model which incorporates only amplitude information well predicted the three populations of lock-in phases. When phase information was removed from the electrosensory stimulus, phase-locking behavior was still observed. These results suggest that phase-locking behavior of Adontosternarchus requires amplitude information but not phase information available in the electrosensory stimulus. [ABSTRACT FROM AUTHOR]

Losses of cone opsin genes are noted in animals that are nocturnal or rely on senses other than vision. We investigated the cone opsin repertoire of night-active South American weakly electric fish. We obtained opsin gene sequences from genomic DNA of 3 gymnotiforms (Eigenmannia virescens, Sternopygus macrurus, Apteronotus albifrons) and the assembled genome of the electric eel (Electrophorus electricus). We identified genes for long-wavelength-sensitive (LWS) and medium-wavelength-sensitive cone opsins (RH2) and rod opsins (RH1). Neither of the 2 short-wavelength-sensitive cone opsin genes were found and are presumed lost. The fact that Electrophorus has a complete repertoire of extraretinal opsin genes and conservation of synteny with the zebrafish (Danio rerio) for genes flanking the 2 short-wavelength-sensitive opsin genes supports the supposition of gene loss. With microspectrophotometry and electroretinograms we observed absorption spectra consistent with RH1 and LWS but not RH2 opsins in the retinal photoreceptors of E. virescens. This profile of opsin genes and their retinal ex-pression is identical to the gymnotiform's sister group, the catfish, which are also nocturnally active and bear ampullary electroreceptors, suggesting that this pattern likely occurred in the common ancestor of gymnotiforms and catfish. Finally, we noted an unusual N-terminal motif lacking a conserved glycosylation consensus site in the RH2 opsin of gymnotiforms, a catfish and a characin (Astyanax mexicanus). Mutations at this site influence rhodopsin trafficking in mammalian photoreceptors and cause retinitis pigmentosa. We speculate that this unusual N terminus may be related to the absence of the RH2 opsin in the cones of gymnotiforms and catfish. [ABSTRACT FROM AUTHOR]