Patterns of lineage diversification in rabbitfishes

Fishes of the tropical Indo-Pacific family Siganidae comprise 28 species, characterized by their body proportions and their colour patterns. A mitochondrial phylogeny of 20 Siganidae species was produced to infer their evolutionary history. Three distinct, major clades were found, that also correspond to the early radiation of the family into three major ecological types: fusiform species that also live in schools on the inshore reef flats (S. canaliculatus, S. fuscescens, S. luridus, S. rivulatus, S. spinus, S. sutor); deep-bodied species including brightly coloured ones whose adults live in pairs on the reef front (S. corallinus, S. doliatus, S. puellus, S. punctatus, S. unimaculatus, S. virgatus, S. vulpinus), and species that live in small schools in mangroves, estuaries and estuarine lakes (S. guttatus, S. javus, S. lineatus, S. randalli, S. vermiculatus); and a third clade including a cosmopolitan species, S. argenteus, the only species of the family known to possess a pelagic, prejuvenile stage and S. woodlandi, a recently described species from New Caledonia and morphologically close to S. argenteus. The partition of the genus into two sub-genera, Lo (erected for S. unimaculatus, S. vulpinus and three related species possessing a tubular snout) and Siganus (all the other species), had no phylogenetic rationale. The present results indicate that the tubular snout, which apparently results from ecological specialization, is a recent acquisition within the deep-body clade. The Western Indian Ocean endemic S. sutor appeared as the sister-species of the Red Sea endemic S. rivulatus within a well-supported subclade that also included S. canaliculatus and S. fuscescens. S. spinus did not appear as sister-species to S. luridus. S. lineatus haplotypes formed a paraphyletic group with S. guttatus, and an early isolation of Maldives S. lineatus was suggested. Unexpectedly, S. randalli did not appear as the sister-species of S. vermiculatus, but its haplotypes instead were embedded within the West Pacific S. lineatus haplogroup, suggesting recent introgression. Among currently-recognized sister-species with parapatric distribution, S. doliatus and S. virgatus haplotypes formed a single, unresolved haplogroup, as did S. unimaculatus and S. vulpinus. The occurrence of two distinct clades within S. fuscescens was confirmed.     

Cytochrome b barcoding, molecular systematics and geographic differentiation in rabbitfishes (Siganidae)

The fish genus Siganus (Siganidae) is widely distributed in the coastal habitats of all the tropical Indo-Pacific, with 28 nominal species recognized so far, based on general morphology and coloration patterns. A mitochondrial phylogeny of 16 Siganidae species, based on the partial nucleotide sequences of the cytochome b gene, was produced. Individual haplotypes of given nominal species generally clustered at the extremity of long branches, thus validating the current taxonomy. However, S. lineatus haplotypes formed a paraphyletic group including S. guttatus, while S. fuscescens haplotypes were apparently splitted in two groups, calling for further investigation. S. woodlandi and S. argenteus formed a monophyletic group, as expected from their close morphological relatedness, although they were separated by a substantial, 14.5-16.3% nucleotide distance. Among eight species sampled from different locations across the Indo-West Pacific, S. argenteus and S. spinus showed the lowest degree of geographic differentiation, a result that correlated well with their extended pelagic larval stage. Fixation index estimates were high in all six other species tested (S. doliatus, S. fuscescens, S. lineatus, S. puellus, S. punctatus, S. vulpinus). The cytochrome b gene fragment chosen here proved useful as a barcode in Siganidae.     

Dactylogyrids (Monogenoidea) parasitizing the gills of spinefoots (Teleostei: Siganidae): proposal of Glyphidohaptor n. gen., with two new species from the Great Barrier Reef, Australia, and G. plectocirra n. comb. from Ras Mohammed National Park, Egypt

Nine species of Siganus (Perciformes: Siganidae) were examined for dactylogyrids (Monogenoidea) from the Red Sea, Egypt; the Great Barrier Reef, Australia; and the South China Sea, China. Species of Tetrancistrum were found on siganids from all 3 localities; Pseudohaliotrema spp. were restricted to siganids from the Great Barrier Reef; and species representing Glyphidohaptor n. gen. were found on siganids from the Red Sea and Great Barrier Reef. Siganus argenteus from the Red Sea and Siganus vulpinus from the Great Barrier Reef were negative for dactylogyrid parasites. Glyphidohaptor n. gen. is proposed for 3 species (2 species new to science) and the new species are described: Glyphidohaptor phractophallus n. sp. from Siganus fuscescens from the Great Barrier Reef; Glyphidohaptor sigani n. sp. from Siganus doliatus (type host), Siganus punctatus, Siganus corallinus, and Siganus lineatus from the Great Barrier Reef; and Glyphidohaptor plectocirra (Paperna, 1972) n. comb. (= Pseudohaliotrema plectocirra Paperna, 1972) from Siganus luridus and Siganus rivulatus from the Red Sea.     

Hybridization between pallid sturgeon Scaphirhynchus albus and shovelnose sturgeon Scaphirhynchus platorynchus

This study found that introgressive hybridization of the pallid sturgeon Scaphirhynchus albus with the common shovelnose sturgeon Scaphirhynchus platorynchus has probably occurred across the range of S. albus. Bayesian clustering found evidence of hybridization in all management units of S. albus. Some individuals were intermediate at both genetic and morphological characters, and some had discordant results. The results support introgressive hybridization throughout much of the range of S. albus, yet individuals consistent with being pure members of each species were detected in all management units. Simulations demonstrated that it would be very difficult to distinguish introgressed individuals from pure specimens after multiple generations of backcrossing with these microsatellite markers. Using hybrid or backcross fish as broodstock could artificially accelerate the loss of unique genetic variation in S. albus. Additional microsatellite loci or additional genetic markers, along with morphological data may be required to ensure that hybrid or backcross fish are not used. Introgressive hybridization requires at least two generations and generation lengths of S. albus are long, perhaps as long as 30 years. The proportion of individuals consistent with introgressive hybrid origins indicates that hybridization between S. albus and S. platorynchus probably has occurred for several generations and is not a recent phenomenon.     

Hybridization between invasive Spartina densiflora (Poaceae) and native S. foliosa in San Francisco Bay, California, USA

Rapid evolution in contemporary time can result when related species, brought together through human-aided introduction, hybridize. The significant evolutionary consequences of post-introduction hybridization range from allopolyploid speciation to extinction of species through genetic amalgamation. Both processes are known to occur in the perennial cordgrass genus, Spartina. Here we report the existence of a third recent Spartina hybridization, discovered in 2002, between introduced S. densiflora and native S. foliosa in San Francisco Bay, California, USA. We used nuclear and chloroplast DNA analysis and nuclear DNA content with chromosome counts to examine plants of morphology intermediate between S. densiflora and S. foliosa in a restored marsh in Marin County, California. We found 32 F(1) diploid hybrids and two triploid plants, all having S. densiflora and S. foliosa as parents; there is also evidence of a genetic contribution of S. alterniflora in some hybrids. None of these hybrids set germinable seed. In 2007 we found a hybrid over 30 miles away in a marsh where both parental species occurred, suggesting hybridization may not be a localized phenomenon. The presence of diploid and triploid hybrids is important because they indicate that several avenues existed that may have given rise to a new allopolyploid species. However, such an event is now unlikely because all hybrids are targets of eradication efforts.     

Functional divergence between morphs of a dwarf chameleon: differential locomotor kinematics in relation to habitat structure

Arboreal lizards are extremely effective at moving in structurally complex habitats, including surfaces of varying diameter and incline. Chameleons exemplify this by exhibiting a number of morphological specializations for moving in these habitats, including the use of prehensile feet and tail to grasp branches. Despite their unique morphology and behaviour, little is known about how locomotor movements vary between species. In addition, some species, such as the Cape Dwarf Chameleon, Bradypodion pumilum, consist of two morphs that differ in ecology, morphology, and behaviour. The two morphs can be found in either closed canopy woodland habitat or relatively open fynbos habitat. The morph that occupies the woodland habitat tends to be larger and utilizes larger diameter perches. Although their ecological and morphological divergence is established, whether this translates into differences in three‐dimensional kinematics of locomotion is not known. Given the potentially strong selective pressures from structurally different habitats, kinematic differences might reveal the functional basis of incipient speciation. We determined that the two morphs diverge significantly in multidimensional kinematic space, and that this occurs for the forelimb and hindlimb independently. These differences outweigh the effects of substrate within each morph, although the differences between morphs were more pronounced on the vertical treatments.     

Microsatellite and morphological analysis of population structure in the parthenogenetic freshwater snail Melanoides tuberculata: insights into the creation of clonal variability

The distribution of variability was studied at various geographical scales in the tropical freshwater snail Melanoides tuberculata, in order to analyse the role of factors shaping this distribution, including the mating system and population dynamics. This parthenogenetic polyploid species reproduces mainly asexually, with males occurring at low frequency. About 800 individuals (38 sites) were sampled from Africa and the Middle East, where the species originated, and from recently colonized habitats in South and Central America, and especially the island of Martinique. We first described variation of general aspects and ornamentation of the shells. This analysis confirms the existence of discrete morphs. Second, individuals were studied at three microsatellite loci, showing that each morph is a genetic clone with some minor variation compatible with models of microsatellite evolution. The genetic analysis also showed much more variation within than between clones. However, two populations from Africa exhibited a large amount of variability, and a mixture of sexual and asexual reproduction might explain these genetic patterns. The worldwide distribution of variability is, therefore, compatible with the African origin of the species, and the introduction of a few clones in other parts of the world. These results also suggest that the distribution of variability in Martinique is influenced by flooding events, and that two morphs from Martinique can be interpreted as hybrids between two pre-existing morphs, based on morphological, genetic and geographical arguments.     

Introgressive hybridization between color morphs in a population of cichlid fishes twelve years after human-induced secondary admixis

In the extremely species-rich haplochromine cichlid fishes of the East African Great Lakes, prezygotic isolation between closely related species is often maintained by color-assortative mating. In 1998, local fisherman working for the ornamental fish trade released different color morphs of the cichlid genus Tropheus into a small harbor basin in the southern part of Lake Tanganyika. This artificial amalgamation of color morphs provides a unique possibility to study mating patterns in cichlids in a natural environment over time. In a precursor study, we analyzed genotypes and phenotypes of almost 500 individuals sampled between 1999 and 2001 and uncovered a marked degree of color-assortative mating, which depended on the level of color pattern dissimilarity between morphs. Twelve years after introduction of nonindigenous morphs, we again sampled Tropheus individuals from the harbor basin and an adjacent, originally pure population and analyzed phenotypes (coloration) and genotypes (mitochondrial control region and 9 microsatellite loci) to assess the current status of the admixed population. Principal component analyses of color score data and population assignment tests demonstrate an increasing level of introgressive hybridization between morphs but also some ongoing color-assortative mating within morphs. The observed mating pattern might have been influenced by fluctuating environmental conditions such as periodic algal blooms or increased sedimentation causing turbid conditions in an otherwise clear lake.     

Space use and genetic structure do not maintain color polymorphism in a species with alternative behavioral strategies

Space use including territoriality and spatial arrangement within a population can reveal important information on the nature, dynamics, and evolutionary maintenance of alternative strategies in color polymorphic species. Despite the prevalence of color polymorphic species as model systems in evolutionary biology, the interaction between space use and genetic structuring of morphs within populations has rarely been examined. Here, we assess the spatial and genetic structure of male throat color morphs within a population of the tawny dragon lizard, Ctenophorus decresii. Male color morphs do not differ in morphology but differ in aggressive and antipredator behaviors as well as androgen levels. Despite these behavioral and endocrine differences, we find that color morphs do not differ in territory size, with their spatial arrangement being essentially random with respect to each other. There were no differences in genetic diversity or relatedness between morphs; however, there was significant, albeit weak, genetic differentiation between morphs, which was unrelated to geographic distance between individuals. Our results indicate potential weak barriers to gene flow between some morphs, potentially due to nonrandom pre‐ or postcopulatory mate choice or postzygotic genetic incompatibilities. However, space use, spatial structure, and nonrandom mating do not appear to be primary mechanisms maintaining color polymorphism in this system, highlighting the complexity and variation in alternative strategies associated with color polymorphism.     

Ongoing Speciation in the Tibetan Plateau Gymnocypris Species Complex

Local adaptation towards divergent ecological conditions often results in genetic differentiation and adaptive phenotypic divergence. To illuminate the ecological distinctiveness of the schizothoracine fish, we studied a Gymnocypris species complex consisting of three morphs distributed across four bodies of water (the Yellow River, Lake Qinghai, the Ganzi River and Lake Keluke) in the Northeast Tibetan Plateau. We used a combination of mitochondrial (16S rRNA and Cyt b) and nuclear (RAG-2) genetic sequences to investigate the phylogeography of these morphs based on a sample of 277 specimens. Analysis of gill rakers allowed for mapping of phenotypic trajectories along the phylogeny. The phylogenetic and morphological analyses showed that the three sparsely rakered morphs were present at two extremes of the phylogenetic tree: the Yellow River morphs were located at the basal phylogenetic split, and the Lake Keluke and Ganzi River morphs at the peak, with the densely rakered Lake Qinghai morphs located between these two extremes. Age estimation further indicated that the sparsely rakered morphs constituted the oldest and youngest lineages, whereas the densely rakered morph was assigned to an intermediate-age lineage. These results are most compatible with the process of evolutionary convergence or reversal. Disruptive natural selection due to divergent habitats and dietary preferences is likely the driving force behind the formation of new morphs, and the similarities between their phenotypes may be attributable to the similarities between their forms of niche tracking associated with food acquisition. This study provides the first genetic evidence for the occurrence of convergence or reversal in the schizothoracine fish of the Tibetan Plateau at small temporal scales.     

Distinct patterns of hybridization across a suture zone in a coral reef fish (Dascyllus trimaculatus)

Hybrid zones are natural laboratories for investigating the dynamics of gene flow, reproductive isolation, and speciation. A predominant marine hybrid (or suture) zone encompasses Christmas Island (CHR) and Cocos (Keeling) Islands (CKE), where 15 different instances of interbreeding between closely related species from Indian and Pacific Oceans have been documented. Here, we report a case of hybridization between genetically differentiated Pacific and Indian Ocean lineages of the three‐spot dascyllus, Dascyllus trimaculatus (Rüppell, 1829). Field observations indicate there are subtle color differences between Pacific and Indian Ocean lineages. Most importantly, population densities of color morphs and genetic analyses (mitochondrial DNA and SNPs obtained via RADSeq) suggest that the pattern of hybridization within the suture zone is not homogeneous. At CHR, both color morphs were present, mitochondrial haplotypes of both lineages were observed, and SNP analyses revealed both pure and hybrid genotypes. Meanwhile, in CKE, the Indian Ocean color morphs were prevalent, only Indian Ocean mitochondrial haplotypes were observed, and SNP analysis showed hybrid individuals with a large proportion (~80%) of their genotypes assigning to the Indian Ocean lineage. We conclude that CHR populations are currently receiving an influx of individuals from both ocean basins, with a greater influence from the Pacific Ocean. In contrast, geographically isolated CKE populations appear to be self‐recruiting and with more influx of individuals from the Indian Ocean. Our research highlights how patterns of hybridization can be different at scales of hundreds of kilometers, due to geographic isolation and the history of interbreeding between lineages.     

Natural hybridization between black spruce and red spruce

Using species-specific random amplified polymorphic DNA (RAPD) markers and morphological characters, natural hybridization between the closely related black spruce Picea mariana (Mill.) B.S.P. and red spruce P. rubens Sarg. was evaluated in natural populations from north-eastern North America. Sampling included populations from both areas of allopatry and also 14 populations from part of the area of sympatry located in the province of Québec and covering several thousands of square kilometres. Classification results from RAPD species-specific markers and from a discriminant function based on morphology were compared. Molecular analysis of the allopatric populations indicated a small amount of interspecific gene leakage with no asymmetric directionality to introgression. A high occurrence of hybrid/introgressant individuals was observed within sympatric populations, suggesting weak reproductive isolation. As expected, the detection of such individuals was more efficient using molecular markers than with morphological traits. The hybrid zone appeared extensive with variable species structure and, in some stands, the main component composed of hybrid/introgressant trees. Implications for the genecology and genetic management of these species are discussed.     

Hybridization between native barbless carp (Cyprinus pellegrini) and introduced common carp (C. carpio) in Xingyun Lake, China

Hybridization with introduced fish species is an important threat to native fish species. Here we investigated hybridization between native barbless carp (Cyprinus pellegrini) and introduced common carp (C. carpio) in Xingyun Lake in the Yunnan-Guizhou plateau of China. A total of 203 individuals of Cyprinus from Xingyun Lake were studied by combination of morphological and genetic analyses. Most individuals were strictly intermediate between the two parental species in morphology, strongly suggesting that extensive hybridization has occurred. Bayesian model-based clustering of the genetic data suggests that there are two distinct genetic groups corresponding to barbless and common carp respectively. Many individuals in the two genetic groups showed intermediate morphology, suggesting that both groups actually contain massively introgressed genes. Only two individuals were identified as barbless carp both morphologically and genetically, hinting that this native species is at risk of genetic extinction in Xingyun Lake.     

Re-evaluation of morphological and chloroplast DNA variation in Juniperus osteosperma Hook and Juniperus occidentalis Torr. Little (Cupressaceae) and their putative hybrids

Relationships between variation in cpDNA and morphology were examined to test the hypothesis of hybridization between Juniperus osteosperma and Juniperus occidentalis. Principal components analysis of 11 taxonomically-important characters distinguished individuals collected from eastern Nevada and Utah from those of southern Oregon. In contrast, many individuals collected from sympatric populations in western Nevada were morphologically intermediate to these two groups. Comparative sequencing of the trnS-trnG intergenic spacer and restriction site analysis of a trnL-trnF PCR product revealed nine haplotypes, and examination of haplotype-morphology associations allowed identification of species-specific genetic markers as well as those that transcend species limits. The confinement of morphological intermediacy and transcendent haplotypes to zones of sympatry, the discovery of haplotypes characteristic of one species in the morphological background of the other, and determination that the character intermediacy encountered is of the type expected under interspecific gene flow are marshaled in support of introgressive hybridization between J. osteosperma and J. occidentalis.     

Discordant patterns of morphological and genetic divergence in the closely related species Schizanthus hookeri and S.?grahamii (Solanaceae)

In this study I have examined the patterns of morphological and genetic differentiation between two species of the Andean genus Schizanthus that differ in their pollination and mating systems. Schizanthus hookeri has a bee pollination syndrome and is strongly dependent on pollinators for seed set. In contrast, S.?grahamii has a hummingbird pollination syndrome and exhibits late autonomous selfing. Southern populations of the latter species have red flowers (reddish morph), while northern populations have yellow (yellowish morph) or pink flowers (pinkish morph). I used two noncoding chloroplast DNA (cpDNA) regions to investigate the genetic affinities between S.?hookeri and the three morphs of S.?grahamii. I also performed intra- and interspecific crosses to assess whether gene flow between species was possible. Phylogenetic analyses supported the existence of two differentiated clades that did not match currently accepted taxonomic classification. Accordingly, genetic distance did not correlate significantly with morphological distance. No fruits were produced from interspecific crosses, and there were no individuals with intermediate morphology that could indicate current and frequent hybridization events between species.?I propose that the discordance between cpDNA data and conventional taxonomy could be explained by parallel evolution, or alternatively, by a very sporadic hybridization.     

Maintenance of genetic and morphological identity in two sibling Syrrhopodon species (Calymperaceae,Bryopsida) despite extensive introgression

Bryophytes are a group of land plants in which the role of hybridization has long been challenged. Using genotyping by sequencing to circumvent the lack of molecular variation at selected loci previously used for phylogeny and morphology, we determine the level of genetic and morphological divergence and reproductive isolation between the sibling Syrrhopodon annotinus and S. simmondsii (Calymperaceae, Bryopsida) that occur in sympatry but in different habitats in lowland Amazonian rainforests. A clear morphological differentiation and a low (0.06), but significant Fst derived from the analysis of 183 single nucleotide polymorphisms were observed between the two species. Conspecific pairs of individuals consistently exhibited higher average kinship coefficients along a gradient of geographic isolation than interspecific pairs. The weak, but significant genetic divergence observed is consistent with growing evidence that ecological specialization can lead to genetic differentiation among bryophyte species. Nevertheless, the spatial genetic structures of the two species were significantly correlated, as evidenced by the significant slope of the Mantel test based on kinship coefficients between pairs of interspecific individuals and the geographic distance separating them. Interspecific pairs of individuals are thus more closely related when they are geographically closer, suggesting that isolation‐by‐distance is stronger than the interspecific reproductive barrier and pointing to interspecific gene flow. We conclude that interspecific introgression, whose role has long been questioned in bryophytes, may take place even in species wherein sporophyte production is scarce due to dioicy, raising the question as to what mechanisms maintain differentiation despite weak reproductive isolation.     

A preliminary study, using multivariate analysis, of a species complex of African house snakes (Boaedon fuliginosus)

systematics of Black and Lined house snakes from the Boaedon fuliginosus species complex were investigated using the limited amount of readily available material. These two forms have recently been regarded both as separate species, and as conspecific morphs. Multivariate and univariate biometrics indicate that in West Africa these two forms are separate species, B. fuliginosus (Black house snake) and B. lineatus (Lined house snake). However, in the rest of Africa these two forms appear to be the conspecific morphs of a single species which is provisionally and tentatively regarded as B. fuliginosus even though it is predominantly striped. There is evidence of "character shift" and this species complex may pose some interesting problems for the concepts of "gene pool cohesion" and "phenetic species".     

Correlated morphological and colour differences among females of the damselfly Ischnura elegans

Abstract 1. The female‐limited colour polymorphic damselfly Ischnura elegans has proven to be an interesting study organism both as an example of female sexual polymorphism, and in the context of the evolution of colour polymorphism, as a model of speciation processes. 2. Previous research suggests the existence of correlations between colour morph and other phenotypic traits, and the different female morphs in I. elegans may be pursuing alternative phenotypically integrated strategies. However, previous research on morphological differences in southern Swedish individuals of this species was only carried out on laboratory‐raised offspring from a single population, leaving open the question of how widespread such differences are. 3. The present study therefore analysed multi‐generational data from 12 populations, investigating morphological differences between the female morphs in the field, differences in the pattern of phenotypic integration between morphs, and quantified selection on morphological traits. 4. It was found that consistent morphological differences indeed existed between the morphs across populations, confirming that the previously observed differences were not simply a laboratory artefact. It was also found, somewhat surprisingly, that despite the existence of sexual dimorphism in body size and shape, patterns of phenotypic integration differed most between the morphs and not between the sexes. Finally, linear selection gradients showed that female morphology affected fecundity differently between the morphs. 5. We discuss the relevance of these results to the male mimicry hypothesis and to the existence of potential ecological differences between the morphs.     

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.