Evidence for Parapatric Speciation in the Mormyrid Fish, Pollimyrus castelnaui (Boulenger, 1911), from the Okavango–Upper Zambezi River Systems: P. marianne sp. nov., Defined by Electric Organ Discharges, Morphology and Genetics

We report on parapatric speciation in the mormyrid fish, Pollimyrus castelnaui (Boulenger, 1911), from the Okavango and the Upper Zambezi River systems. We recognise samples from the Zambezi River as a distinct species, P. marianne, displaying an eastern phenotype of electric organ discharge (EOD) waveform (Type 3) that is distinct from the western EOD phenotype (Type 1) observed in P. castelnaui samples from the neighbouring Okavango. Samples from the geographically intermediate Kwando/Linyanti River (a tributary of the Zambezi that is also intermittently connected to the Okavango) presented a more variable third EOD phenotype (Type 2). In 13 out of 14 morphological characters studied, the Zambezi River samples differed significantly from P. castelnaui. Morphologically and in EOD characters, the Kwando/Linyanti fish are distinct from both P. castelnaui and P. marianne. Sequence analysis of the mitochondrial cytochrome b gene unambiguously reveals that specimens from the Zambezi River System form a well supported taxon which clearly differs from P. castelnaui from the Okavango (1.5–2.5% sequence divergence). Within specimens from the Kwando–Zambezi System some geographic differentiation can be detected (nucleotide substitutions up to 0.6%); but groups cannot be resolved with certainty. Significant allozyme differences were found between the Okavango and all other EOD types from the Upper Zambezi System, and, within the Zambezi System, between the Kwando (Type 2) and Zambezi (Type 3) individuals. The low Wright's fixation index values, the lack of fixed allele differences, and small genetic distances provide little evidence for speciation between groups within the Zambezi System, but moderate to great fixation index values and significant allele frequency differences were observed between the Okavango and the other fishes. It is concluded that within the Zambezi System, differentiation between Kwando/Linyanti and Zambezi populations (as revealed by morphology and EOD waveform comparisons) is so recent that substantial genetic (allozyme and mitochondrial sequence) differences could not have evolved, or were not detected.     

The Southern Churchill, Petrocephalus Wesselsi, a New Species of Mormyrid from South Africa defined by Electric Organ Discharges, Genetics, and Morphology

East African and south African churchills (Petrocephalus, Mormyridae) were synonymised in 1959 to become members of a single species of subcontinental, southern African distribution, Petrocephalus catostoma (Günther, 1866). By comparison with the type material for P. catostoma from the Ruvuma River and P. stuhlmanni from the Ruvu River, both of East African origin, we confirm the South African form of churchill to represent a new species, P. wesselsi, ranging from the northern Limpopo and Incomati systems south to the Pongola River (Natal) as its southern limit. We also compared churchills from the Sabie River (25° S, South Africa, Incomati system) with churchills from the Upper Zambezi River (17° S, Namibia), using electric organ discharges (EODs) and morphology. The duration of an EOD pulse of the South African form (N = 39; 943.2±S.E. 18.82µs) is, on average, more than twice that of the Upper Zambezi form (N = 37; 436.6±15.1µs), and the amplitude of the second head-positive phase (P2 phase relative to P1 = 1) significantly weaker (0.133 ± 0.0005 vs. 0.472 ± 0.002 for Upper Zambezi males, 0.363 ± 0.03 for Upper Zambezi females). In contrast to the Upper Zambezi form, the EOD of the South African form exhibits no difference between the sexes. Fish from the two origins differ significantly in 11 out of 14 anatomical characters studied, confirming molecular genetic differentiation on the species level.     

Comparing the fish assemblages and food‐web structures of large floodplain rivers

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

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

Validation of annulus formation in sagittal otoliths of three tropical cichlids Oreochromis andersonii (Castelnau, 1861), Oreochromis macrochir (Boulenger, 1912), and Coptodon rendalli (Boulenger, 1896)

The periodicity of growth zone formation in sagittal otoliths of Oreochromis andersonii, Oreochromis macrochir and Coptodon rendalli from the Upper Zambezi region, southern Africa, was validated to test the hypothesis that a single growth zone is deposited annually. Edge analysis was carried out on 754 O. andersonii (54–492 mm TL), 601 O. macrochir (83–438 mm TL) and 622 C. rendalli (45–379 mm TL) caught between August 2010 and August 2011 in Lake Liambezi and the Kavango, Kwando and Upper Zambezi rivers. This was corroborated for O. andersonii (n = 17) and C. rendalli (n = 16) using a fluorochrome marking experiment conducted in a large earthen pond (625 m², 1.5 m deep) adjacent to the Kavango River. Both methods demonstrated that growth zone deposition occurred annually in juvenile and adult fishes. For edge analysis, the highest proportion of otoliths with an opaque margin was observed in summer from October to December. The fluorochrome marking experiment, however, revealed that opaque zone deposition took place in winter between June and September when water temperatures were lowest (17°C), indicating a lag between opaque zone deposition and detection on the otolith edge.     

Observations on the electric organ discharge of two skate species (Chondrichthyes: Rajidae) and its relationship to behaviour

Synopsis The electric organ discharge (EOD) of the little skate,Raja erinacea and winter skate,R. ocellata was recorded both from isolated individuals and from small groups using methods that allowed for the identification of individuals producing EODs. Pulse duration, train lengh, frequency, and pulse patterns are characterized and correlated with behaviour. The two species,R. erinacea andR. ocellata, were found to have characteristically different EOD pulse durations of 70 ms and 217 ms respectively. Isolated skates rarely discharged whereas groups of skates were found to discharge regularly. The EOD was evoked by tactile prodding, physical contact with other skates and electrical stimulation. Skates also discharged reflexively in response to an artificially induced head-positive DC stimulus, sine wave and monopolar square pulses. During approach and contact, skates responded to each other with interacting EOD displays. EOD interaction and pulse duration differences between other species suggest a possible intra-specific communication function of the EOD inRaja.     

Genetic diversity of two Southern African cichlids (Oreochromis andersonii and O. macrochir) in the Zambezi and Congo River basins

Oreochromis andersonii and O. macrochir are two important cichlid species native to Southern Africa. We describe in this paper their genetic population structure in the Upper Zambezi River, Kafue River, and Lake Bangweulu representing part of the Congo and Zambezi River Basins. Microsatellite genetic markers were employed to analyse the genetic population structure of the two species using 177 tissue samples. The average allele richness of O. andersonii was higher in the Zambezi River (10.500) than in the Kafue River (9.583) though not statistically different. For O. macrochir, it was highest in the Zambezi River (11.170) followed by the Kafue River (9.781) and least in Lake Bangweulu (7.067) and their differences were significant. The gene diversity indices; gene diversity (hs), observed heterozygosity (H_O), expected heterozygosity (H_E) were high and similar in O. andersonii populations. However, in O. macrochir H_E was significantly lower in Lake Bangweulu (0.678) compared to the Kafue River (0.799) and Zambezi River (0.802) populations. Population differentiation estimated by R_ST and D_EST revealed high differentiation in both species (R_ST = 0.598, D_EST = 0.777 for O. andersonii; R_ST = 0.379, D_EST = 0.710 for O. macrochir). The highest source of variation was among populations (84.71%) for O. andersonii and within populations (67.09%) for O. macrochir. Comparisons of population pairs revealed a close genetic similarity between the Zambezi River and Lake Bangweulu populations of O. macrochir. Bottlenecks were observed in both species using the Two-Phase Model (T.P.M.) indicative of a recent genetic loss or reduction in effective population size. Though our results indicate that the populations of both species still maintain sufficiently high levels of genetic diversity in the sampled areas, the bottlenecks observed are a source of concern. We recommend a more robust study of genetic diversity of these species in all sections of these river systems and that some key conservation sites should be identified to protect the gene pool of these native species.     

Hormonal control of sex differences in the electric organ discharge (EOD) of mormyrid fishes

Summary Field studies have demonstrated that several species of mormyrid fish from Gabon, West Africa have a sex difference in the pulse-like waveform of their Electric Organ Discharge (EOD). Administration of androgen hormones (testosterone or dihydrotestosterone) to a female or juvenile can induce the EOD typical of a sexually mature male. Data for two such species —Brienomyrus brachyistius (triphasic) andStomatorhinus corneti — are presented, showing that transformation of a female's or juvenile's EOD to a male-like EOD involves a 2–3 fold increase in EOD duration and a downward shift in the peak frequency of the EOD's power spectrum (as determined by Fourier analysis). ForBrienomyrus brachyistius (triphasic), estradiol can also induce changes in the EOD waveform, although not as dramatic as that for androgens. Changes in EOD duration and power spectra are often accompanied by an alteration of the wave-shape or morphology of the EOD pulse, i.e., the relative amplitude of its peaks and the presence of inflection points in its negative and positive phases.A third species,Hippopotamyrus batesii (triphasic), not previously known to have an EOD sex difference, also responds to testosterone treatment with an increase in EOD duration. Preliminary field data indicate this species may have a sexual dimorphism in its EOD, suggesting that the response to a steroid hormone may be an indicator of a sex difference in a species' EOD waveform. Such findings are discussed in relation to the affects of steroids on vertebrate neurons and muscle, and the evolution of electric communication systems.Abbreviations DHT 5-dihydrotestosterone - EOD electric organ discharge - F mature female - HTI interval between peaks (H and T) in EOD's first derivative - IF juvenile female - IM juvenile male - M mature male - PPW peak frequency of power spectrum     

Phylogeography and speciation in the <Emphasis Type="Italic">Pseudocrenilabrus philander</Emphasis> species complex in Zambian Rivers

Haplochromine cichlids form the most species-rich lineage of cichlid fishes that both colonized almost all river systems in Africa and radiated to species flocks in several East African lakes. The enormous diversity of lakes is contrasted by a relatively poor albeit biogeographically clearly structured species diversity in rivers. The present study analyzed the genetic structure and phylogeographic history of species and populations of the genus Pseudocrenilabrus in Zambian rivers that span two major African drainage systems, the Congo- and the Zambezi-system. The mtDNA phylogeny identifies four major lineages, three of which occur in the Congo-system and one in the Zambezi system. Two of the Congo-clades (Lake Mweru and Lunzua River) comprise distinct albeit yet undescribed species, while the fish of the third Congo-drainage clade (Chambeshi River and Bangweulu swamps), together with the fish of the Zambezi clade (Zambezi and Kafue River) are assigned to Pseudocrenilabrus philander. Concerning the intraspecific genetic diversity observed in the sampled rivers, most populations are highly uniform in comparison to lacustrine haplochromines, suggesting severe founder effects and/or bottlenecking during their history. Two bursts of diversification are reflected in the structure of the linearized tree. The first locates at about 3.9% mean sequence divergence and points to an almost simultaneous colonization of the sampled river systems. Subsequent regional diversification (with about 1% mean sequence divergence) occurred contemporaneously within the Kafue River and the Zambezi River. The clear-cut genetic biogeographic structure points to the dominance of geographic speciation in this lineage of riverine cichlid fishes, contrasting the importance of in situ diversification observed in lake cichlids.     

Integrative taxonomy reveals hidden diversity in the southern African darters genus Nannocharax Günther 1867 (Characiformes: Distichodontidae)

The present study explored the diversity of Nannocharax within southern Africa by implementing three species delimitation methods for a data set consisting of 37 mitochondrial cytochrome oxidase subunit I sequences. Two unilocus coalescent methods, the General Mixed Yule Coalescent (GMYC) and the Bayesian implementation of the Poisson Tree Processes (bPTP), and a genetic distance method, the Automatic Barcode Gap Discovery (ABGD), were applied. Both GMYC and bPTP delimited the same operational taxonomic units (OTUs), revealing a higher diversity for the genus in the region than previously recognised, whereas the ABGD failed to delimit the same candidate species. All methods delimited two species groups, and these are supported based on colouration patterning and morphology; the Nannocharax multifasciatus and the Nannocharax macropterus species groups and the delimited OTUs were assigned to each. Two putative new species were identified, Nannocharax cf. lineostriatus “Okavango” from the Okavango River in Angola and N. cf. lineostriatus “Kwanza” from the Kwanza River system in Angola. The distribution of Nannocharax dageti was confirmed for the Upper Zambezi and extended to the Okavango system, and an identification key for the southern Africa Nannocharax species is provided.     

Genetic and phylogenetic evidence for misidentification of Vibrio species within the Harveyi clade

Aim: This report describes the use of a six‐gene multi‐locus sequence analysis (MLSA) to correctly identify Vibrio strains of the Harveyi clade. Methods and Results: Vibrio isolates were characterized using a six housekeeping gene MLSA. The study provided evidence supporting: (i) a substantial number of reference strains maintained within commercial culture collections are misidentified taxonomically at the species level; (ii) two V. alginolyticus subclades retain species‐level divergence; and (iii) V. communis and V. owensii likely are the same species. Conclusion: A significant number (n = 10) of Harveyi clade Vibrio strains have been inaccurately identified, including evidence that V. communis and V. owensii strains, two recently discovered species assigned to the Harveyi clade, comprise a single species. Significance and Impact of the study: As Harveyi clade vibrios have an enormous impact on human and aquatic animal health, it is of paramount importance to identify members of the Harveyi clade correctly.     

Parasites of fishes in the recently inundated ephemeral Lake Liambezi,Namibia

Lake Liambezi forms the periodic connection between the upper Zambezi, Kwando and Okavango rivers. A full parasitological assessment was conducted on 86 fish, representing 14 species in six families sampled in August 2011. Parasite diversity was low and dominated by species with complex life cycles involving intermediate hosts. Most prevalent were larval nematodes (Contracaecum sp.) infecting 12 and Trypanasoma sp. infecting nine of the 14 host species. The intra-erythrocytic parasite Babesiosoma mariae was found in the blood of Coptodon rendalli and Oreochromis andersonii with prevalence of 50% and 60%, respectively. The host-specific monogenean Annulotrema hepseti was recorded only from H. cuvieri with a prevalence of 100%. Notable absences were the copepod and branchiuran parasites that have direct lifecycles and usually occur in high prevalence and abundance in the region. Because parasites with direct life cycles can only be transported into the lake on the host fish, their absence suggests limited immigration of infected fishes into the lake. This suggests that internal recruitment dominates over immigration in the fish population dynamics in Lake Liambezi.     

Comparative ecology of Serranochromis species (Teleostei: Cichlidae) in the Upper Zambezi River floodplain

The Upper Zambezi River drainage of Zambia contains six Serrunochromis species belonging to the largemouth, piscivorous subgenus Serranochromis and three smallmouth, invertebrate-feeding species of the subgenus Sargochromis. A survey of the Barotse floodplain and associateddrainages revealed that two species, S. (S.) longimanus (Boulenger) and S. (S.) thumbergi (Castelnau), were uncommon on the floodplain. S. thumbergi appeared to be more abundant in the Kafue, Chobe, and Kabompo rivers than the Upper Zambezi proper. Adults of the three largest species (i.e. S. (S) robustus (Günther), S. (S.) altus (Winemiller & Kelso-Winemiller), and S. (Sar.) giurdi (Pellegrin)) inhabit the main channel of the Zambezi River during low water, whereas most adults of the other four species (i.e., S. (S.) macrocephalus (Boulenger), S. (S.)ungusticeps (Boulenger), S. (Sar.) codringtoni (Boulenger), and S. (Sar.) carlottae (Boulenger)) were collected from lagoons, canals and backwaters. Comparisons of population size structure and gonad condition during the falling water (May-August) and low water (September- December) seasons indicated that the seven common Serranochromis initiate spawning near the end of the low water period. Examination of gonad condition during the pre-spawning period indicated that Serrunochromis attain maturity at the following sizes: S. robustus, males (275-300 mm s.l .), females (250-275 mm); S. macrocephalus, both sexes (150-200 mm); S. altus, males (275-325 mm), females (250-300 mm);S. angusticeps, males (250-275 mm), females (175-225 mm); S. giardi, males (275-300 mm), females (225-250 mm); S. codringtoni, males (150 mm), females (125-150 mm); S. carlottue, males (150-175 mm), females (100-125 mm). Age was estimated from scale annuli comprised of closely spaced circuli that corresponded to the period of rising water (December- March). S. robustus, S. altus, S. angusticeps, and S. giurdi matured at approximately 3 years, and S. macrocephalus, S. codringtoni, and S. curlottae matured at approximately 2 years in the Barotse floodplain. The average numbers of mature oocytes per female were as follows: S. robustus- 1165; S. macrocephalus-810; S. altus-920; S. angusticeps-627; S. codringtoni-580; S. carlottue-362. Fecundity exhibited a weak positive relationship with female size. Analysis of stomach contents showed that interspecific diet overlap was low among both immature and mature size classes of Serrunochromis. Adult size classes of S. robustus fed primarily on small Synodontis catfishes, S. mucrocephalus fed mostly on mormyrid and cichlid fishes, S. altus specialized on mormyrids, and S. angusticeps fed primarily on small cyprinid and characid fishes. The three members of the subgenus Sargochromis consumed mostly invertebrates. Molluscs were the most important item in the diet of S. giardi, and aquatic insects and fish scales were predominant in the diet of S. carlottae. Larger size classes of S. codringtoni consumed large amounts of seeds (Graminaceae). If habitat and diet are considered together, niche separation among Upper Zambezi Serrunochromis species appears to be nearly complete.     

Vegetation-environment relations of a Middle Zambezi floodplain

Detrended correspondence analysis was used to study the relationships between environmental factors and the species composition of vegetation on Zambezi River alluvium downstream of the Kariba hydroelectric dam. Grass, sedge and woody species were recorded in 73 stands in Mana Pools National Park, Zimbabwe. Grass and sedge species composition was related to the soil moisture regime, as indexed by soil texture and flooding frequency. The first woody plant ordination axis was related to a stand development gradient; Acacia albida was a pioneer species on lowlying sandbanks and the woody species richness of stands increased with their height above the Zambezi River. Two-way indicator species analysis identified 7 vegetation types which could be separated on the basis of their topsoil texture and flooding frequency. The types were: sandbanks; young A. albida woodland; A. albida woodland; A. albida dominated mixed woodland; mixed riverine woodland with understory; mixed riverine woodland; and grassland on clay soils.     

Two new species of Gelidium (Rhodophyta, Gelidiales), Gelidium tenuifolium and Gelidium koshikianum, from Japan

Two new marine red algae, Gelidium tenuifolium sp. nov. and Gelidium koshikianum sp. nov. (Gelidiales, Gelidiaceae) are described from Japan. Gelidium tenuifolium with large‐sized thalli (up to 30 cm tall) is distinguished from other species with such thalIi by the production of wide, flattened and thin branches (up to 2 mm wide and 60–80 urn thick), the presence of an apical depression and simple determinate branches. Gelidium koshikianum with middle‐sized thalli (5–8 cm tall) is distinguished from other species with such thalli by having wide axes (up to 2.5 mm wide) and short (2.0–3.2 mm), unbranched, second‐ and third‐order branches issuing at short intervals (0.8–1.4 mm). In phylogenetic analyses of rbcL sequences, four Gelidium species that are chiefly distributed in Japan including G. tenuifolium were clustered together with 99% bootstrap value (Japanese Gelidium‐complex clade). Gelidium linoides Kützing came to the position of the sister group to G. tenuifolium with 99% bootstrap value. There were four substitutions (0.3% divergence) between G. linoides and G. tenuifolium sequences. Gelidium koshikianum and Gelidium allanii Chapman were clustered together with 100% bootstrap value and they came to the position of the sister group to the Japanese GeIidium‐complex clade with 83% bootstrap value. There were six substitutions (0.4% divergence) between G. koshikianum and G. allanii sequences.     

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

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