Phylogenomics of Bony-Tongue Fishes (Osteoglossomorpha) Shed Light on the Craniofacial Evolution and Biogeography of the Weakly Electric Clade (Mormyridae).

Bonytongues (Osteoglossomorpha) constitute an ancient clade of teleost fishes distributed in freshwater habitats throughout the world. The group includes well-known species such as arowanas, featherbacks, pirarucus, and the weakly electric fishes in the family Mormyridae. Their disjunct distribution, extreme morphologies, and electrolocating capabilities (Gymnarchidae and Mormyridae) have attracted much scientific interest, but a comprehensive phylogenetic framework for comparative analysis is missing, especially for the species-rich family Mormyridae. Of particular interest are disparate craniofacial morphologies among mormyrids which might constitute an exceptional model system to study convergent evolution. We present a phylogenomic analysis based on 546 exons of 179 species (out of 260), 28 out of 29 genera, and all six families of extant bonytongues. Based on a recent reassessment of the fossil record of osteoglossomorphs, we inferred dates of divergence among transcontinental clades and the major groups. The estimated ages of divergence among extant taxa (e.g., Osteoglossomorpha, Osteoglossiformes, and Mormyroidea) are older than previous reports, but most of the divergence dates obtained for clades on separate continents are too young to be explained by simple vicariance hypotheses. Biogeographic analysis of mormyrids indicates that their high species diversity in the Congo Basin is a consequence of range reductions of previously widespread ancestors and that the highest diversity of craniofacial morphologies among mormyrids originated in this basin. Special emphasis on a taxon-rich representation for mormyrids revealed pervasive misalignment between our phylogenomic results and mormyrid taxonomy due to repeated instances of convergence for extreme craniofacial morphologies. Estimation of ancestral phenotypes revealed contingent evolution of snout elongation and unique projections from the lower jaw to form the distinctive Schnauzenorgan. Synthesis of comparative analyses suggests that the remarkable craniofacial morphologies of mormyrids evolved convergently due to niche partitioning, likely enabled by interactions between their exclusive morphological and electrosensory adaptations. [Africa; ancestral state estimation; diversity; exon capture; freshwater fishes; Phylogenomics.].

Comparison of ideal mask-based speech enhancement algorithms for speech mixed with white noise at low mixture signal-to-noise ratios.

The literature shows that the intelligibility of noisy speech can be improved by applying an ideal binary or soft gain mask in the time-frequency domain for signal-to-noise ratios (SNRs) between -10 and +10 dB. In this study, two mask-based algorithms are compared when applied to speech mixed with white Gaussian noise (WGN) at lower SNRs, that is, SNRs from -29 to -5 dB. These comprise an Ideal Binary Mask (IBM) with a Local Criterion (LC) set to 0 dB and an Ideal Ratio Mask (IRM). The performance of three intrusive Short-Time Objective Intelligibility (STOI) variants-STOI, STOI+, and Extended Short-Time Objective Intelligibility (ESTOI)-is compared with that of other monaural intelligibility metrics that can be used before and after mask-based processing. The results show that IRMs can be used to obtain near maximal speech intelligibility (>90% for sentence material) even at very low mixture SNRs, while IBMs with LC = 0 provide limited intelligibility gains for SNR < -14 dB. It is also shown that, unlike STOI, STOI+ and ESTOI are suitable metrics for speech mixed with WGN at low SNRs and processed by IBMs with LC = 0 even when speech is high-pass filtered to flatten the spectral tilt before masking.

Intelligibility prediction for speech mixed with white Gaussian noise at low signal-to-noise ratios.

The effect of additive white Gaussian noise and high-pass filtering on speech intelligibility at signal-to-noise ratios (SNRs) from -26 to 0 dB was evaluated using British English talkers and normal hearing listeners. SNRs below -10 dB were considered as they are relevant to speech security applications. Eight objective metrics were assessed: short-time objective intelligibility (STOI), a proposed variant termed STOI+, extended short-time objective intelligibility (ESTOI), normalised covariance metric (NCM), normalised subband envelope correlation metric (NSEC), two metrics derived from the coherence speech intelligibility index (CSII), and an envelope-based regression method speech transmission index (STI). For speech and noise mixtures associated with intelligibility scores ranging from 0% to 98%, STOI+ performed at least as well as other metrics and, under some conditions, better than STOI, ESTOI, STI, NSEC, CSIIMid, and CSIIHigh. Both STOI+ and NCM were associated with relatively low prediction error and bias for intelligibility prediction at SNRs from -26 to 0 dB. STI performed least well in terms of correlation with intelligibility scores, prediction error, bias, and reliability. Logistic regression modeling demonstrated that high-pass filtering, which increases the proportion of high to low frequency energy, was detrimental to intelligibility for SNRs between -5 and -17 dB inclusive.

Finite difference time domain modelling of a point-excited elastic plate radiating into an acoustic cavity.

Finite difference time domain (FDTD) models are developed to solve the vibroacoustic problem of a thin elastic plate undergoing point force excitation and radiating into an acoustic cavity. Vibroacoustic modelling using FDTD can be computationally expensive because structure-borne sound wavespeeds are relatively high and a fine spatial resolution is often required. In this paper a scaling approach is proposed and validated to overcome this problem through modifications to the geometry and physical properties. This allows much larger time steps to be used in the model which significantly reduces the computation time. Additional reductions in computation time are achieved by introducing an alternative approach to model the boundaries between the air and the solid media. Experimental validation is carried out using a thin metal plate inside a small reverberant room. The agreement between FDTD and measurements confirms the validity of both approaches as well as the FDTD implementation of a thin plate as a three-dimensional solid that can support multiple wave types. Below the lowest room mode, there are large spatial variations in the sound field within the cavity due to the radiating plate; this indicates the importance of having a validated FDTD model for low-frequency vibroacoustic problems.

Thresholds of information leakage for speech security outside meeting rooms.

This paper describes an approach to provide speech security outside meeting rooms where a covert listener might attempt to extract confidential information. Decision-based experiments are used to establish a relationship between an objective measurement of the Speech Transmission Index (STI) and a subjective assessment relating to the threshold of information leakage. This threshold is defined for a specific percentage of English words that are identifiable with a maximum safe vocal effort (e.g., "normal" speech) used by the meeting participants. The results demonstrate that it is possible to quantify an offset that links STI with a specific threshold of information leakage which describes the percentage of words identified. The offsets for male talkers are shown to be approximately 10 dB larger than for female talkers. Hence for speech security it is possible to determine offsets for the threshold of information leakage using male talkers as the "worst case scenario." To define a suitable threshold of information leakage, the results show that a robust definition can be based upon 1%, 2%, or 5% of words identified. For these percentages, results are presented for offset values corresponding to different STI values in a range from 0.1 to 0.3.

Prediction of high-frequency vibration transmission across coupled, periodic ribbed plates by incorporating tunneling mechanisms.

Prediction of structure-borne sound transmission on built-up structures at audio frequencies is well-suited to Statistical Energy Analysis (SEA) although the inclusion of periodic ribbed plates presents challenges. This paper considers an approach using Advanced SEA (ASEA) that can incorporate tunneling mechanisms within a statistical approach. The coupled plates used for the investigation form an L-junction comprising a periodic ribbed plate with symmetric ribs and an isotropic homogeneous plate. Experimental SEA (ESEA) is carried out with input data from Finite Element Methods (FEM). This indicates that indirect coupling is significant at high frequencies where bays on the periodic ribbed plate can be treated as individual subsystems. SEA using coupling loss factors from wave theory leads to significant underestimates in the energy of the bays when the isotropic homogeneous plate is excited. This is due to the absence of tunneling mechanisms in the SEA model. In contrast, ASEA shows close agreement with FEM and laboratory measurements. The errors incurred with SEA rapidly increase as the bays become more distant from the source subsystem. ASEA provides significantly more accurate predictions by accounting for the spatial filtering that leads to non-diffuse vibration fields on these more distant bays.

Differential expression of genes and proteins between electric organ and skeletal muscle in the mormyrid electric fish Brienomyrus brachyistius.

Electric organs (EOs) have evolved independently in vertebrates six times from skeletal muscle (SM). The transcriptional changes accompanying this developmental transformation are not presently well understood. Mormyrids and gymnotiforms are two highly convergent groups of weakly electric fish that have independently evolved EOs: while much is known about development and gene expression in gymnotiforms, very little is known about development and gene expression in mormyrids. This lack of data limits prospects for comparative work. We report here on the characterization of 28 differentially expressed genes between SM and EO tissues in the mormyrid Brienomyrus brachyistius, which were identified using suppressive subtractive hybridization (SSH). Forward and reverse SSH was performed on tissue samples of EO and SM resulting in one cDNA library enriched with mRNAs expressed in EO, and a second library representing mRNAs unique to SM. Nineteen expressed sequence tags (ESTs) were identified in EO and nine were identified in SM using BLAST searching of Danio rerio sequences available in NCBI databases. We confirmed differential expression of all 28 ESTs using RT-PCR. In EO, these ESTs represent four classes of proteins: (1) ion pumps, including the α- and ß-subunits of Na(+)/K(+)-ATPase, and a plasma membrane Ca(2+)-ATPase; (2) Ca(2+)-binding protein S100, several parvalbumin paralogs, calcyclin-binding protein and neurogranin; (3) sarcomeric proteins troponin I, myosin heavy chain and actin-related protein complex subunit 3 (Arcp3); and (4) the transcription factors enhancer of rudimentary homolog (ERH) and myocyte enhancer factor 2A (MEF2A). Immunohistochemistry and western blotting were used to demonstrate the translation of seven proteins (myosin heavy chain, Na(+)/K(+)-ATPase, plasma membrane Ca(2+)-ATPase, MEF2, troponin and parvalbumin) and their cellular localization in EO and SM. Our findings suggest that mormyrids express several paralogs of muscle-specific genes and the proteins they encode in EOs, unlike gymnotiforms, which may post-transcriptionally repress several sarcomeric proteins. In spite of the similarity in the physiology and function of EOs in mormyrids and gymnotiforms, this study indicates that the mechanisms of development in the two groups may be considerably different.

Mitogenome recovered from a 19 th Century holotype by shotgun sequencing supplies a generic name for an orphaned clade of African weakly electric fishes (Osteoglossomorpha, Mormyridae).

Heteromormyrus Steindachner, 1866, a genus of Mormyridae (Teleostei: Osteoglossomorpha), has been monotypic since the description of Heteromormyruspauciradiatus (Steindacher, 1866) from a single specimen. No type locality other than "Angola" was given and almost no specimens have been subsequently identified to this species. In order to investigate the relationship of this taxon to fresh specimens collected in Angola and elsewhere, whole genome paired-end sequencing of DNA extracted from the holotype specimen of Heteromormyruspauciradiatus was performed and a nearly complete mitogenome assembled from the sequences obtained. Comparison of cytochrome oxidase I and cytochrome b sequences from this mitogenome to sequences from recently collected material reveal that Heteromormyruspauciradiatus is closely related to specimens identified as Hippopotamyrusansorgii (Boulenger, 1905), Hippopotamyrusszaboi Kramer, van der Bank & Wink, 2004, Hippopotamyruslongilateralis Kramer & Swartz, 2010, as well as to several undescribed forms from subequatorial Africa collectively referred to in the literature as the "Hippopotamyrusansorgii species complex" and colloquially known as "slender stonebashers." Previous molecular phylogenetic work has shown that these species are not close relatives of Hippopotamyruscastor Pappenheim, 1906, the type species of genus Hippopotamyrus Pappenheim, 1906 from Cameroon, and are thus misclassified. Hippopotamyrusansorgii species complex taxa and another species shown to have been misclassified, Paramormyropstavernei (Poll, 1972), are placed in genus Heteromormyrus and one genetic lineage from the Kwanza and Lucala rivers of Angola are identified as conspecific Heteromormyruspauciradiatus. Three additional new combinations and a synonymy in Mormyridae are introduced. The morphological characteristics and geographical distribution of the genus Heteromormyrus are reviewed. The electric organ discharges (EODs) of Heteromormyrus species are to be treated in a separate study.

Signal variation and its morphological correlates in Paramormyrops kingsleyae provide insight into the evolution of electrogenic signal diversity in mormyrid electric fish.

We describe patterns of geographic variation in electric signal waveforms among populations of the mormyrid electric fish species Paramormyrops kingsleyae. This analysis includes study of electric organs and electric organ discharge (EOD) signals from 553 specimens collected from 12 localities in Gabon, West-Central Africa from 1998 to 2009. We measured time, slope, and voltage values from nine defined EOD "landmarks" and determined peak spectral frequencies from each waveform; these data were subjected to principal components analysis. The majority of variation in EODs is explained by two factors: the first related to EOD duration, the second related to the magnitude of the weak head-negative pre-potential, P0. Both factors varied clinally across Gabon. EODs are shorter in eastern Gabon and longer in western Gabon. Peak P0 is slightly larger in northern Gabon and smaller in southern Gabon. P0 in the EOD is due to the presence of penetrating-stalked (Pa) electrocytes in the electric organ while absence is due to the presence of non-penetrating stalked electrocytes (NPp). Across Gabon, the majority of P. kingsleyae populations surveyed have only individuals with P0-present EODs and Pa electrocytes. We discovered two geographically distinct populations, isolated from others by barriers to migration, where all individuals have P0-absent EODs with NPp electrocytes. At two sites along a boundary between P0-absent and P0-present populations, P0-absent and P0-present individuals were found in sympatry; specimens collected there had electric organs of intermediate morphology. This pattern of geographic variation in EODs is considered in the context of current phylogenetic work. Multiple independent paedomorphic losses of penetrating stalked electrocytes have occurred within five Paramormyrops species and seven genera of mormyrids. We suggest that this key anatomical feature in EOD signal evolution may be under a simple mechanism of genetic control, and may be easily influenced by selection or drift throughout the evolutionary history of mormyrids.

On the efficacy of spatial sampling using manual scanning paths to determine the spatial average sound pressure level in rooms.

In architectural acoustics, noise control and environmental noise, there are often steady-state signals for which it is necessary to measure the spatial average, sound pressure level inside rooms. This requires using fixed microphone positions, mechanical scanning devices, or manual scanning. In comparison with mechanical scanning devices, the human body allows manual scanning to trace out complex geometrical paths in three-dimensional space. To determine the efficacy of manual scanning paths in terms of an equivalent number of uncorrelated samples, an analytical approach is solved numerically. The benchmark used to assess these paths is a minimum of five uncorrelated fixed microphone positions at frequencies above 200 Hz. For paths involving an operator walking across the room, potential problems exist with walking noise and non-uniform scanning speeds. Hence, paths are considered based on a fixed standing position or rotation of the body about a fixed point. In empty rooms, it is shown that a circle, helix, or cylindrical-type path satisfy the benchmark requirement with the latter two paths being highly efficient at generating large number of uncorrelated samples. In furnished rooms where there is limited space for the operator to move, an efficient path comprises three semicircles with 45°-60° separations.

Sexual signal evolution outpaces ecological divergence during electric fish species radiation.

Natural selection arising from resource competition and environmental heterogeneity can drive adaptive radiation. Ecological opportunity facilitates this process, resulting in rapid divergence of ecological traits in many celebrated radiations. In other cases, sexual selection is thought to fuel divergence in mating signals ahead of ecological divergence. Comparing divergence rates between naturally and sexually selected traits can offer insights into processes underlying species radiations, but to date such comparisons have been largely qualitative. Here, we quantitatively compare divergence rates for four traits in African mormyrid fishes, which use an electrical communication system with few extrinsic constraints on divergence. We demonstrate rapid signal evolution in the Paramormyrops species flock compared to divergence in morphology, size, and trophic ecology. This disparity in the tempo of trait evolution suggests that sexual selection is an important early driver of species radiation in these mormyrids. We also found slight divergence in ecological traits among closely related species, consistent with a supporting role for natural selection in Paramormyrops diversification. Our results highlight the potential for sexual selection to drive explosive signal divergence when innovations in communication open new opportunities in signal space, suggesting that opportunity can catalyze species radiations through sexual selection, as well as natural selection.

Attitudes to Noise Inside Dwellings in Three Megacities: Seoul, London, and São Paulo.

This study investigated people's attitudes towards noise inside their homes. Online questionnaire surveys were conducted in Seoul, London, and São Paulo. The questionnaire was designed to assess annoyance caused by noise from neighbours and environmental noise (transportation). Information was also collected on situational, personal, and socio-demographic variables. Respondents that were more annoyed by outdoor noise inside their dwelling reported higher neighbour noise annoyance. In Seoul, neighbour noise was found to be more annoying than outdoor noise, and those with higher noise sensitivity reported higher annoyance towards neighbour noise. However, neighbour noise and outdoor noise was found to be equally annoying in London and São Paulo. For neighbour noise, the average percentage of respondents hearing structure-borne sources compared to airborne sources differed in each city. Most neighbour noise sources in São Paulo gave rise to higher annoyance ratings than Seoul and London. Education and income levels had a limited effect on annoyance and coping strategy. Annoyance with indoor noise from neighbours was found to have stronger relationships with cognitive and behavioural coping strategies than outdoor noise annoyance.

Petrocephalus of Odzala offer insights into evolutionary patterns of signal diversification in the Mormyridae, a family of weakly electrogenic fishes from Africa.

Electric signals of mormyrid fishes have recently been described from several regions of Africa. Members of the Mormyridae produce weak electric organ discharges (EODs) as part of a specialized electrosensory communication and orientation system. Sympatric species often express distinctive EODs, which may contribute to species recognition during mate choice in some lineages. Striking examples of interspecific EOD variation within assemblages have been reported for two monophyletic radiations: the Paramormyrops of Gabon and the Campylomormyrus of Lower Congo. Here, we describe a speciose assemblage of Petrocephalus in the Lékoli River system of Odzala National Park, Republic of Congo. This widespread genus comprises the subfamily (Petrocephalinae) that is the sister group to all other mormyrids (Mormyrinae). Eleven Petrocephalus species were collected in Odzala, five of which are not described taxonomically. We quantify EOD variation within this assemblage and show that all eleven species produce EOD waveforms of brief duration (species means range from 144 to 663mus) compared to many other mormyrids. We also present reconstructed phylogenetic relationships among species based on cytochrome b sequences. Discovery of the Odzala assemblage greatly increases the number of Petrocephalus species for which EODs and DNA sequence data are available, permitting a first qualitative comparison between mormyrid subfamilies of the divergence patterns that have been described within lineages. We find that the Petrocephalus assemblage in Odzala is not a monophyletic radiation. Genetic divergence among Petrocephalus species often appears higher than among Paramormyrops or Campylomormyrus species. In contrast, results of this study and others suggest that Petrocephalus may generally exhibit less interspecific EOD divergence, as well as smaller sex differences in EOD waveforms, compared to Paramormyrops and Campylomormyrus. We discuss possible causes and consequences of EOD diversification patterns observed within mormyrid subfamilies as a framework for future comparative studies of signal evolution using this emerging model system.

Comparison of high-intensity sound and mechanical vibration for cleaning porous titanium cylinders fabricated using selective laser melting.

Orthopedic components, such as the acetabular cup in total hip joint replacement, can be fabricated using porous metals, such as titanium, and a number of processes, such as selective laser melting. The issue of how to effectively remove loose powder from the pores (residual powder) of such components has not been addressed in the literature. In this work, we investigated the feasibility of two processes, acoustic cleaning using high-intensity sound inside acoustic horns and mechanical vibration, to remove residual titanium powder from selective laser melting-fabricated cylinders. With acoustic cleaning, the amount of residual powder removed was not influenced by either the fundamental frequency of the horn used (75 vs. 230 Hz) or, for a given horn, the number of soundings (between 1 and 20). With mechanical vibration, the amount of residual powder removed was not influenced by the application time (10 vs. 20 s). Acoustic cleaning was found to be more reliable and effective in removal of residual powder than cleaning with mechanical vibration. It is concluded that acoustic cleaning using high-intensity sound has significant potential for use in the final preparation stages of porous metal orthopedic components. © 2015 Wiley Periodicals, Inc. J Biomed Mater Res Part B: Appl Biomater, 105B: 117-123, 2017.

Cryptomyrus: a new genus of Mormyridae (Teleostei, Osteoglossomorpha) with two new species from Gabon, West-Central Africa.

We use mitochondrial and nuclear sequence data to show that three weakly electric mormyrid fish specimens collected at three widely separated localities in Gabon, Africa over a 13-year period represent an unrecognized lineage within the subfamily Mormyrinae and determine its phylogenetic position with respect to other taxa. We describe these three specimens as a new genus containing two new species. Cryptomyrus, new genus, is readily distinguished from all other mormyrid genera by a combination of features of squamation, morphometrics, and dental attributes. Cryptomyrus ogoouensis, new species, is differentiated from its single congener, Cryptomyrus ona, new species, by the possession of an anal-fin origin located well in advance of the dorsal fin, a narrow caudal peduncle and caudal-fin lobes nearly as long as the peduncle. In Cryptomyrus ona, the anal-fin origin is located only slightly in advance of the dorsal fin, the caudal peduncle is deep and the caudal-fin lobes considerably shorter than the peduncle. Continued discovery of new taxa within the "Lower Guinea Clade" of Mormyridae highlights the incompleteness of our knowledge of fish diversity in West-Central Africa. We present a revised key to the mormyrid genera of Lower Guinea.

RésuméNous avons démontré avec des marqueurs mitochondriaux et nucléaires que trois spécimens de poissons mormyridés faiblement électriques, collectés à trois localités au Gabon sur une période de 13 ans, forment une lignée inconnue au sein de la sous famille des Mormyrinae dont nous avons déterminé la position phylogénétique. Nous décrivons un nouveau genre et deux nouvelles espèces pour ces trois spécimens. Cryptomyrus, nouveau genre, se distingue des autres genres de mormyridés par une combinaison de caractères de l'écaillure, morphométriques, et dentaires. Cryptomyrus ogoouensis, nouvelle espèce, se distingue de sa seule espèce congénérique par sa nageoire anale qui commence nettement en avant de sa nageoire dorsale, son pédoncule caudal gracile et les lobes de sa nageoire caudale pratiquement aussi longs que son pédoncule caudal. Chez Cryptomyrus ona, nouvelle espèce, la nageoire anale commence légèrement en avant de l'origine de la nageoire dorsale, le pédoncule caudal est large, et les lobes de la nageoire caudale sont plus courts que le pédoncule caudal. La découverte ininterrompue de nouveaux taxa dans le «clade de basse Guinée¼ des Mormyridae illustre notre connaissance incomplète de la diversité des poissons de l'ouest de l'Afrique centrale. Nous présentons une clé révisée des genres de Mormyridae de basse Guinée.

Vibrotactile Presentation of Musical Notes to the Glabrous Skin for Adults with Normal Hearing or a Hearing Impairment: Thresholds, Dynamic Range and High-Frequency Perception.

Presentation of music as vibration to the skin has the potential to facilitate interaction between musicians with hearing impairments and other musicians during group performance. Vibrotactile thresholds have been determined to assess the potential for vibrotactile presentation of music to the glabrous skin of the fingertip, forefoot and heel. No significant differences were found between the thresholds for sinusoids representing notes between C1 and C6 when presented to the fingertip of participants with normal hearing and with a severe or profound hearing loss. For participants with normal hearing, thresholds for notes between C1 and C6 showed the characteristic U-shape curve for the fingertip, but not for the forefoot and heel. Compared to the fingertip, the forefoot had lower thresholds between C1 and C3, and the heel had lower thresholds between C1 and G2; this is attributed to spatial summation from the Pacinian receptors over the larger contactor area used for the forefoot and heel. Participants with normal hearing assessed the perception of high-frequency vibration using 1s sinusoids presented to the fingertip and were found to be more aware of transient vibration at the beginning and/or end of notes between G4 and C6 when stimuli were presented 10dB above threshold, rather than at threshold. An average of 94% of these participants reported feeling continuous vibration between G4 and G5 with stimuli presented 10dB above threshold. Based on the experimental findings and consideration of health effects relating to vibration exposure, a suitable range of notes for vibrotactile presentation of music is identified as being from C1 to G5. This is more limited than for human hearing but the fundamental frequencies of the human voice, and the notes played by many instruments, lie within it. However, the dynamic range might require compression to avoid the negative effects of amplitude on pitch perception.

Multiple cases of striking genetic similarity between alternate electric fish signal morphs in sympatry.

Striking trait polymorphisms are worthy of study in natural populations because they can often shed light on processes of phenotypic divergence and specialization, adaptive evolution, and (in some cases) the early stages of speciation. We examined patterns of genetic variation within and between populations of mormyrid fishes that are morphologically cryptic in sympatry but produce alternate types of electric organ discharge (EOD). Other species in a large group containing a clade of these morphologically cryptic EOD types produce stereotyped, species-typical EOD waveforms thought to function in mate recognition. First, for six populations from Gabon's Brienomyrus species flock, we confirm that forms of electric fish that exhibit distinctive morphologies and unique EOD waveforms (i.e., good reference species) are reproductively isolated from coexisting congeners. These sympatric species deviate from genetic panmixia across five microsatellite loci. Given this result, we examined three focal pairs of syntopic and morphologically cryptic EOD waveform types that are notable exceptions to the pattern of robust genetic partitioning among unique waveform classes within assemblages. These exceptional pairs constitute a monophyletic group within the Brienomyrus flock known as the magnostipes complex. One member of each pair (type I) produces a head-negative EOD, while the other member (either type II or type III, depending on location) produces a longer duration EOD differing in waveform from type I. We show that signal development in these pairs begins with juveniles of all magnostipes-complex morphs emitting head-positive EODs resembling those of type II adults. Divergence of EOD waveforms occurs with growth such that there are two discrete and fixed signal types in morphologically indistinguishable adults at each of several localities. Strong microsatellite partitioning between allopatric samples of any of these morphologically cryptic signal types suggests that geographically isolated populations are genetically decoupled from one another. By contrast, sympatric morphs appear genetically identical across microsatellite loci in Mouvanga Creek and the Okano River and only very weakly diverged, if at all, in the Ivindo River. Our results for the magnostipes complex fail to detect species boundaries between the focal morphs and are, instead, fully consistent with the existence of relatively stable signal dimorphisms at each of several different localities. No mechanism for the maintenance of this electrical polymorphism is suggested by the known natural history of the magnostipes complex. Despite a lack of evidence for genetic differentiation, the possibility of incipient sympatric speciation between morphs (especially type I and type II within the Ivindo River) merits further testing due to behavioral and neurobiological lines of evidence implying a general role for stereotyped EOD waveforms in species recognition. We discuss alternative hypotheses concerning the origins, stability, and evolutionary significance of these intriguing electrical morphs in light of geographical patterns of population structure and signal variation.

Discovery and phylogenetic analysis of a riverine species flock of African electric fishes (Mormyridae: Teleostei).

The evolution of species-specific mate recognition signals is of particular interest within speciose monophyletic groups with restricted distributions (known as "species flocks"). However, the explosive nature of speciation in these clades makes difficult the reconstruction of their phylogenetic history. Here we describe a species flock of riverine mormyrid fishes from west-central Africa in which electric signals may play a role in the reproductive isolation of sympatric species. In our recent field collections, totaling more than 1400 specimens from many localities, we recognize 38 forms that are distinct in their morphologies and electric organ discharge (EOD) characteristics. Of these 38, only four clearly correspond to described species. Here we treat these forms as operational taxonomic units (OTUs) in a phylogenetic analysis of cytochrome b sequence data from a sample of 86 specimens. We examined support in the molecular data for the monophyly of these 38 OTUs considered together, the monophyly of each phenotypically delimited OTU considered individually, and for relationships among OTUs congruent with those inferred from the distribution of morphological and EOD character states. Trees obtained by both maximum-parsimony and maximum-likelihood analyses, rooted with sequence data from outgroup taxa, provide evidence for the monophyly of these 38 OTUs with respect to other mormyrid fishes. The small genetic distances between many distinct forms suggest their recent divergence. However, in many instances the cytochrome b tree topology fails to support the monophyly of individual OTUs and close relationships between OTUs that are similar in morphology and EOD characteristics. In other cases, individuals from distinct OTUs share identical or nearly identical haplotypes. Close examination of these cases suggests that unnatural OTU definition is not the sole cause of this pattern, and we infer an incongruence between the mitochondrial gene tree and the organismal phylogeny caused by incomplete mitochondrial lineage sorting and/ or introgression across forms. The apparently rapid diversification in this clade of riverine electric fishes and the problems associated with recovering a meaningful species-level phylogeny from mitochondrial data parallel findings in other species flocks. Selection on EOD waveforms as mate recognition signals may be involved in the radiation of these fishes. This is the first description of a freshwater fish species flock from a riverine, as opposed to a lacustrine, environment.

AFLPs resolve phylogeny and reveal mitochondrial introgression within a species flock of African electric fish (Mormyroidea: Teleostei).

Estimating species phylogeny from a single gene tree can be especially problematic for studies of species flocks in which diversification has been rapid. Here we compare a phylogenetic hypothesis derived from cytochrome b (cyt b) sequences with another based on amplified fragment length polymorphisms (AFLP) for 60 specimens of a monophyletic riverine species flock of mormyrid electric fishes collected in Gabon, west-central Africa. We analyze the aligned cyt b sequences by Wagner parsimony and AFLP data generated from 10 primer combinations using neighbor-joining from a Nei-Li distance matrix, Wagner parsimony, and Dollo parsimony. The different analysis methods yield AFLP tree topologies with few conflicting nodes. Recovered basal relationships in the group are similar between cyt b and AFLP analyses, but differ substantially at many of the more derived nodes. More of the clades recovered with the AFLP characters are consistent with the morphological characters used to designate operational taxonomic units in this group. These results support our hypothesis that the mitochondrial gene tree differs from the overall species phylogeny due at least in part to mitochondrial introgession among lineages. Mapping the two forms of electric organ found in this group onto the AFLP tree suggests that posteriorly innervated electrocytes with nonpenetrating stalks have independently evolved from anteriorly innervated, penetrating-stalk electrocytes at least three times.