Shaw, P W; Turner, G F; Idid, M R; Robinson, R L; Carvalho, G R
Allopatric processes of speciation have routinely been presented to explain the extraordinary radiation of the East African Great Lakes cichlid fish species flocks. The 21 or more species of pelagic cichlids within the Lake Malawi flock appear to have lake-wide distributions that challenge such a concept. Data from six microsatellite DNA loci indicate single, panmictic populations across the lake of three Diplotaxodon species. Levels of variability at these loci suggest that populations have been large and stable. Mitochondrial DNA sequence data (872 bp of control region + 981 bp of the NADH-2) from 90 species, representing all major clades within the Lake Malawi flock, indicate reciprocal monophyly of the pelagic clade. We suggest that these data support a hypothesis that speciation in sympatry is more plausible (and widespread) within the cichlid species flocks than previously thought. PMID:11413643
Smith, Adam R; van Staaden, Moira J; Carleton, Karen L
Although the cichlids of Lake Malawi are an important model system for the study of sensory evolution and sexual selection, the evolutionary processes linking these two phenomena remain unclear. Prior works have proposed that evolutionary divergence is driven by sensory drive, particularly as it applies to the visual system. While evidence suggests that sensory drive has played a role in the speciation of Lake Victoria cichlids, the findings from several lines of research on cichlids of Lake Malawi are not consistent with the primary tenets of this hypothesis. More specifically, three observations make the sensory drive model implausible in Malawi: (i) a lack of environmental constraint due to a broad and intense ambient light spectrum in species rich littoral habitats, (ii) pronounced variation in receiver sensory characteristics, and (iii) pronounced variability in male courtship signal characteristics. In the following work, we synthesize the results from recent studies to draw attention to the importance of sensory variation in cichlid evolution and speciation, and we suggest possible avenues of future research.
Kocher, T D; Conroy, J A; McKaye, K R; Stauffer, J R
The species flocks of cichlid fishes in the lakes of East Africa are the most spectacular example of adaptive radiation among living vertebrates. Similar highly derived morphologies are found among species in different lakes. These similarities have been variously interpreted either as evidence for migration of ancestral species between the lakes, or of striking convergence of morphology. To distinguish among these competing hypotheses we sequenced a portion of the mitochondrial DNA control region from six pairs of morphologically similar taxa from Lakes Malawi and Tanganyika. Our results clearly indicate a separate origin of these morphologies in the two lakes. They also suggest that the Tanganyikan radiation is relatively old, and that the species flocks in lakes Malawi and Victoria are derived from one of the ancient lineages found in Lake Tanganyika. These findings have important implications for understanding the pattern of morphological and behavioral evolution in these fish.
Smith, Adam R; D'Annunzio, Lindsay; Smith, Abbi E; Sharma, Anit; Hofmann, Christopher M; Marshall, N J; Carleton, Karen L
The expression of cone opsin genes is a primary determinant of the characteristics of colour vision. Interspecific variation in opsin expression is common in African cichlids. It is correlated with foraging among cichlids from Lake Malawi, and with ambient light environment among cichlids from Lake Victoria. In this study, we tested whether gene expression varied within species such that it might be important in contributing to divergence. We hypothesized that light attenuation with depth would be correlated with predictable changes in gene expression in Lake Malawi, and that this variation would tune visual sensitivities to match the ambient light environment. We observed significant differences in cone opsin expression in three different comparisons among populations of the same species. Higher LWS expression was found in shallow versus deep Copadichromis eucinostomus. In Metriaclima zebra, individuals from Zimbawe Rock expressed significantly more SWS2B than those from Thumbi West Island, although these locales have similar ambient light environments. Finally, Tropheops gracilior from deeper water had significantly more variation in expression than their shallow counterparts. These results support that gene expression varies significantly between populations of the same species. Surprisingly, these results could not be explained by predicted visual performance as models predicted that differential expression patterns did not confer sensitivity advantages at different depths. This suggested that expression variation did not confer a local sensitivity advantage. Therefore, our findings were contrary to a primary requirement of the sensory bias hypothesis. As such, other explanations for intraspecific gene expression variation need to be tested.
Ono, H; O'hUigin, C; Tichy, H; Klein, J
Lake Malawi in eastern Africa harbors > 500 endemic species of cichlid fishes, all of which are believed to have emerged from a single founding population in the past 2 Myr. Molecular characterization of differences among the species could provide important information about the nature of speciation in the period of adaptive radiation. Because of the close relationship, however, molecular variation among the species has been difficult to ascertain. In this communication, we provide evidence for extensive differences, in major-histocompatibility-complex (Mhc) class II genes, between two related species, Pseudotropheus zebra and Melanochromis auratus. We used specific primers to amplify and sequence intron 1 and exon 2 of the class II genes from 18 individuals. Although we found 20 different sequences among the 42 that we produced, there was not a single sequence shared by the two species. Thus the study suggests that different cichlid species of Lake Malawi have different profiles of class II alleles, presumably because the polymorphism present in the ancestral founding population segregated differentially into the various species. These results make Mhc genes an important tool for elucidating speciation.
Parry, Juliet W L; Carleton, Karen L; Spady, Tyrone; Carboo, Aba; Hunt, David M; Bowmaker, James K
Cichlid fish of the East African Rift Lakes are renowned for their diversity and offer a unique opportunity to study adaptive changes in the visual system in rapidly evolving species flocks. Since color plays a significant role in mate choice, differences in visual sensitivities could greatly influence and even drive speciation of cichlids. Lake Malawi cichlids inhabiting rock and sand habitats have significantly different cone spectral sensitivities. By combining microspectrophotometry (MSP) of isolated cones, sequencing of opsin genes, and spectral analysis of recombinant pigments, we have established the cone complements of four species of Malawi cichlids. MSP demonstrated that each of these species predominately expresses three cone pigments, although these differ between species to give three spectrally different cone complements. In addition, rare populations of spectrally distinct cones were found. In total, seven spectral classes were identified. This was confirmed by opsin gene sequencing, expression, and in vitro reconstitution. The genes represent the four major classes of cone opsin genes that diverged early in vertebrate evolution. All four species possess a long-wave-sensitive (LWS), three spectrally distinct green-sensitive (RH2), a blue-sensitive (SWS2A), a violet-sensitive (SWS2B), and an ultraviolet-sensitive (SWS1) opsin. However, African cichlids determine their spectral sensitivity by differential expression of primarily only three of the seven available cone opsin genes. Phylogenetic analysis suggests that all percomorph fish have similar potential.
Clark, Frances E; Conte, Matthew A; Ferreira-Bravo, Irani A; Poletto, Andreia B; Martins, Cesar; Kocher, Thomas D
B chromosomes are extra chromosomes found in many species of plants, animals, and fungi. B chromosomes often manipulate common cellular processes to increase their frequency, sometimes to the detriment of organismal fitness. Here, we characterize B chromosomes in several species of Lake Malawi cichlid fish. Whole genome sequencing of Metriaclima zebra "Boadzulu" individuals revealed blocks of sequence with unusually high sequence coverage, indicative of increased copy number of those sequences. These regions of high sequence coverage were found only in females. SNPs unique to the high copy number sequences permitted the design of specific amplification primers. These primers amplified fragments only in Metriaclima lombardoi individuals that carried a cytologically identified B chromosome (B-carriers), indicating these extra copies are located on the B chromosome. These same primers were used to identify B-carrying individuals in additional species from Lake Malawi. Across 7 species, a total of 43 B-carriers were identified among 323 females. B-carriers were exclusively female; no B chromosomes were observed in the 317 males surveyed from these species. Quantitative analysis of the copy number variation of B-specific sequence blocks suggests that B-carriers possess a single B chromosome, consistent with previous karyotyping of M. lombardoi A single B chromosome in B-carriers is consistent with 2 potential drive mechanisms: one involving nondisjunction and preferential segregation in a mitotic division prior to the germ-line, and the other involving preferential segregation during meiosis I.
Schwalbe, Margot A B; Webb, Jacqueline F
The adaptive radiations of African cichlids resulted in a diversity of feeding morphologies and strategies, but the role of sensory biology in prey detection and feeding ecology remains largely unexplored. Two endemic Lake Malawi cichlid genera, Tramitichromis and Aulonocara, feed on benthic invertebrates, but differ in lateral line morphology (narrow and widened lateral line canals, respectively) and foraging strategy. The hypothesis that they use their lateral line systems differently was tested by looking at the relative contribution of the lateral line system and vision in prey detection by Tramitichromis sp. and comparing results to those from a complementary study using Aulonocara stuartgranti (Schwalbe et al., 2012). First, behavioral trials were used to assess the ability of Tramitichromis sp. to detect live (mobile) and dead (immobile) benthic prey under light and dark conditions. Second, trials were run before, immediately after, and several weeks after chemical ablation of the lateral line system to determine its role in feeding behavior. Results show that Tramitichromis sp. is a visual predator that neither locates prey in the dark nor depends on lateral line input for prey detection and is thus distinct from A. stuartgranti, which uses its lateral line or a combination of vision and lateral line to detect prey depending on light condition. Investigating how functionally distinctive differences in sensory morphology are correlated with feeding behavior in the laboratory and determining the role of sensory systems in feeding ecology will provide insights into how sensory capabilities may contribute to trophic niche segregation.
O'Quin, Claire T; Drilea, Alexi C; Roberts, Reade B; Kocher, Thomas D
Pigmentation patterns are one of the most recognizable forms of phenotypic diversity and an important component of organismal fitness. While much progress has been made in understanding the genes controlling pigmentation in model systems, many questions remain about the genetic basis of pigment traits observed in nature. Lake Malawi cichlid fishes are known for their diversity of male pigmentation patterns, which have been shaped by sexual selection. To begin the process of identifying the genes underlying this diversity, we quantified the number of pigment cells on the body and fins of two species of the genus Metriaclima and their hybrids. We then used the Castle-Wright equation to estimate that differences in individual pigmentation traits between these species are controlled by one to four genes each. Different pigmentation traits are highly correlated in the F(2) , suggesting shared developmental pathways and genetic pleiotropy. Melanophore and xanthophore traits fall on opposite ends of the first principal component axis of the F(2) phenotypes, suggesting a tradeoff during the development of these two pigment cell types.
Roberts, Reade B; Moore, Emily C; Kocher, Thomas D
Despite long-standing interest in the evolution and maintenance of discrete phenotypic polymorphisms, the molecular genetic basis of such polymorphism in the wild is largely unknown. Female sex-associated blotched colour polymorphisms found in cichlids of Lake Malawi, East Africa, represent a highly successful polymorphic phenotype, found and maintained in four genera across the geographic expanse of the lake. Previously, we identified an association with an allelic variant of the paired-box transcription factor gene pax7a and blotched colour morphs in Lake Malawi cichlid fishes. Although a diverse range of blotched phenotypes are present in Lake Malawi cichlid species, they all appeared to result from an allele of pax7a that produces increased levels of transcript. Here, we examine the developmental and genetic basis of variation among blotched morphs. First, we confirm that pax7a-associated blotch morphs result primarily from modulation of melanophore development and survival. From laboratory crosses and natural population studies, we identify at least three alleles of pax7a associated with discrete subtypes of blotched morphs, in addition to the ancestral pax7a allele. Genotypes at pax7a support initial evolution of a novel pax7a allele to produce the blotched class of morphs, followed by subsequent evolution of that pax7a blotched allele to produce additional alleles associated with discrete colour morphs. Variant alleles of pax7a produce different levels of pax7a transcript, correlating with pigmentation phenotype at the cellular level. This naturally selected allelic series should serve as a case study for understanding the molecular genetic control of pax7a expression and the evolution of sex-associated alleles.
Ivory, Sarah J; Blome, Margaret W; King, John W; McGlue, Michael M; Cole, Julia E; Cohen, Andrew S
Long paleoecological records are critical for understanding evolutionary responses to environmental forcing and unparalleled tools for elucidating the mechanisms that lead to the development of regions of high biodiversity. We use a 1.2-My record from Lake Malawi, a textbook example of biological diversification, to document how climate and tectonics have driven ecosystem and evolutionary dynamics. Before ∼800 ka, Lake Malawi was much shallower than today, with higher frequency but much lower amplitude water-level and oxygenation changes. Since ∼800 ka, the lake has experienced much larger environmental fluctuations, best explained by a punctuated, tectonically driven rise in its outlet location and level. Following the reorganization of the basin, a change in the pacing of hydroclimate variability associated with the Mid-Pleistocene Transition resulted in hydrologic change dominated by precession rather than the high-latitude teleconnections recorded elsewhere. During this time, extended, deep lake phases have abruptly alternated with times of extreme aridity and ecosystem variability. Repeated crossings of hydroclimatic thresholds within the lake system were critical for establishing the rhythm of diversification, hybridization, and extinction that dominate the modern system. The chronology of these changes closely matches both the timing and pattern of phylogenetic history inferred independently for the lake's extraordinary array of cichlid fish species, suggesting a direct link between environmental and evolutionary dynamics.
Schwalbe, Margot A B; Webb, Jacqueline F
Two sand-dwelling cichlids from Lake Malawi (Aulonocara stuartgranti, Tramitichromis sp.) that feed on benthic invertebrates, but have different lateral line phenotypes, use lateral line and/or visual cues to detect prey under light versus dark conditions. The current study examined how ecologically relevant variation in light intensity [0-800 lux (lx)] influences detection of prey (mobile, immobile) in each species by analyzing six behavioral parameters. Both species fed at light intensities ≥1 lx and trends in behavior among light intensities were informative. However, prey type and/or time of day (but not light intensity) predicted all four parameters analyzed with generalized linear mixed models in A. stuartgranti, whereas the interaction of light intensity and time of day predicted three of these parameters in Tramitichromis sp. Data suggest that the critical light intensity is 1-12 lx for both species, that the integration of visual and lateral line input explains differences in detection of mobile and immobile prey and behavioral changes at the transition from 1 to 0 lx in A. stuartgranti, and that Tramitichromis sp. likely uses binocular vision to locate prey. Differences in the sensory biology of species that exploit similar prey will have important implications for the trophic ecology of African cichlid fishes.
Arnegard, M. E.; Markert, J. A.; Danley, P. D.; Stauffer, J. R.; Ambali, A. J.; Kocher, T. D.
Extremely fine-scale genetic partitioning has recently been detected among populations of Lake Malawi's rock-dwelling cichlids through the study of microsatellite loci. Understanding the mechanisms of genetic differentiation that operate in this rapidly speciating group requires further investigation of the geographic patterns of gene flow and the congruence between morphological and genetic divergence. In pursuit of this goal, genetic variation at four microsatellite loci and variation in male breeding coloration were examined in several populations of Labeotropheus fuelleborni from southern Lake Malawi. Significant genetic differentiation exists among populations (overall FST = 0.063; p = 0.0002). While migration appears unrestricted within continuous rocky patches, deep waters and sandy bays more than 2 km wide act as strong barriers to gene flow. Dispersal of L. fuelleborni appears to follow a stepping-stone model in which the distribution of habitats often constrains migration to one dimension. It is hypothesized that clinal colour variation in the study area has resulted from the secondary contact of divergent lineages, although reproductive isolation between colour variants is not apparent. Relative to shoreline populations, reduced levels of gene flow among populations inhabiting isolated, deep-water islands provides greater opportunities for drift, adaptation to local conditions, or sexual selection to effect genetic differentiation in this species.
Cetron, M S; Chitsulo, L; Sullivan, J J; Pilcher, J; Wilson, M; Noh, J; Tsang, V C; Hightower, A W; Addiss, D G
Schistosomiasis is a parasitic infection caused by trematodes. Humans are infected through skin contact with free-swimming cercariae which develop in freshwater snails. Schistosomiasis has been endemic to Malawi for several decades, but the open waters and shores of Lake Malawi have long been thought to be risk-free with regard to schistosomiasis transmission. However, in 1992, two US Peace Corps volunteers developed central nervous system schistosomiasis due to infection with Schistosoma haematobium following recreational water exposure at Cape Maclear on Lake Malawi. In light of these infections, a cross-sectional survey of resident expatriates and visitors to Malawi was subsequently conducted during March-April 1993 to determine the transmission potential and risk for acquiring schistosomiasis in the lake. 305 US citizens and 650 non-US foreign nationals participated in the study. Serological evidence of current or past schistosome infection was identified in 303 subjects. Indeed, seroprevalence was 32% among expatriates whose freshwater exposure was limited to Lake Malawi; S. haematobium antibodies were found in 135 of 141 seropositive specimens. The risk of seropositivity increased with the number of freshwater exposures at Lake Malawi resorts. While many resort areas in the southwestern lake region posed a significant risk, Cape Maclear was the location most strongly associated with seropositivity. Schistosome-infected Bulinus globosus, the snail vector of S. haematobium in Malawi, were found at Cape Maclear and other locations along the lakeshore.
Brzozowski, Frances; Roscoe, Jennifer; Parsons, Kevin; Albertson, Craig
East African cichlids are renowned for their propensity to radiate, and variation in color patterns accounts for much of endemic cichlid diversity. Sexual dimorphism in color among cichlid species likely represents the outcome of different selective regimes acting on each sex, and is a classic example of sexual conflict. It is generally assumed that this conflict has been mitigated through the evolution of sex-linked color polymorphisms. Here, we propose that the evolution of sex-specific differences in levels of color trait integration may represent an additional mechanism through which sexual conflict has been resolved in this group. Specifically, we predict: (1) that general patterns of integration are influenced by early developmental events and thus conserved across sexes and (2) that male color is less integrated than females, and thus more evolvable in terms of producing an elaborate palette (i.e., in response to sexual selection), whereas female color is more integrated, facilitating wholesale shifts in color for background matching (i.e., in response to natural selection for crypsis). We tested these hypotheses using an F(2) design to compare the segregation of male and female color patterns. Both exploratory methods and hypothesis-driven analyses of integration demonstrate that the covariance structure of color traits in males and females is distinct, and that males are significantly less integrated than females. We suggest that the ability of species to promote different levels, and to a lesser extent patterns, of phenotypic integration between males and females may have contributed to the evolutionary success of this group.
Shirai, Kazumasa; Inomata, Nobuyuki; Mizoiri, Shinji; Aibara, Mitsuto; Terai, Yohey; Okada, Norihiro; Tachida, Hidenori
When a population size is reduced, genetic drift may fix slightly deleterious mutations, and an increase in nonsynonymous substitution is expected. It has been suggested that past aridity has seriously affected and decreased the populations of cichlid fishes in Lake Victoria, while geographical studies have shown that the water levels in Lake Tanganyika and Lake Malawi have remained fairly constant. The comparably stable environments in the latter two lakes might have kept the populations of cichlid fishes large enough to remove slightly deleterious mutations. The difference in the stability of cichlid fish population sizes between Lake Victoria and the Lakes Tanganyika and Malawi is expected to have caused differences in the nonsynonymous/synonymous ratio, ω (=dN/dS), of the evolutionary rate. Here, we estimated ω and compared it between the cichlids of the three lakes for 13 mitochondrial protein-coding genes using maximum likelihood methods. We found that the lineages of the cichlids in Lake Victoria had a significantly higher ω for several mitochondrial loci. Moreover, positive selection was indicated for several codons in the mtDNA of the Lake Victoria cichlid lineage. Our results indicate that both adaptive and slightly deleterious molecular evolution has taken place in the Lake Victoria cichlids' mtDNA genes, whose nonsynonymous sites are generally conserved.
Nagl, Sandra; Tichy, Herbert; Mayer, Werner E.; Takahata, Naoyuki; Klein, Jan
Phylogenetic trees for groups of closely related species often have different topologies, depending on the genes used. One explanation for the discordant topologies is the persistence of polymorphisms through the speciation phase, followed by differential fixation of alleles in the resulting species. The existence of transspecies polymorphisms has been documented for alleles maintained by balancing selection but not for neutral alleles. In the present study, transspecific persistence of neutral polymorphisms was tested in the endemic haplochromine species flock of Lake Victoria cichlid fish. Putative noncoding region polymorphisms were identified at four randomly selected nuclear loci and tested on a collection of 12 Lake Victoria species and their putative riverine ancestors. At all loci, the same polymorphism was found to be present in nearly all the tested species, both lacustrine and riverine. Different polymorphisms at these loci were found in cichlids of other East African lakes (Malawi and Tanganyika). The Lake Victoria polymorphisms must have therefore arisen after the flocks now inhabiting the three great lakes diverged from one another, but before the riverine ancestors of the Lake Victoria flock colonized the Lake. Calculations based on the mtDNA clock suggest that the polymorphisms have persisted for about 1.4 million years. To maintain neutral polymorphisms for such a long time, the population size must have remained large throughout the entire period. PMID:9826684
Nagl, S; Tichy, H; Mayer, W E; Takahata, N; Klein, J
Phylogenetic trees for groups of closely related species often have different topologies, depending on the genes used. One explanation for the discordant topologies is the persistence of polymorphisms through the speciation phase, followed by differential fixation of alleles in the resulting species. The existence of transspecies polymorphisms has been documented for alleles maintained by balancing selection but not for neutral alleles. In the present study, transspecific persistence of neutral polymorphisms was tested in the endemic haplochromine species flock of Lake Victoria cichlid fish. Putative noncoding region polymorphisms were identified at four randomly selected nuclear loci and tested on a collection of 12 Lake Victoria species and their putative riverine ancestors. At all loci, the same polymorphism was found to be present in nearly all the tested species, both lacustrine and riverine. Different polymorphisms at these loci were found in cichlids of other East African lakes (Malawi and Tanganyika). The Lake Victoria polymorphisms must have therefore arisen after the flocks now inhabiting the three great lakes diverged from one another, but before the riverine ancestors of the Lake Victoria flock colonized the Lake. Calculations based on the mtDNA clock suggest that the polymorphisms have persisted for about 1.4 million years. To maintain neutral polymorphisms for such a long time, the population size must have remained large throughout the entire period.
Sturmbauer, C; Meyer, A
Of the three cichlid species flocks in eastern Africa, Lake Tanganyika harbors the oldest species assemblage, which is also the most diverse morphologically and behaviorally. For 12 species (20 individuals) of 12 genera of the tribe Ectodini, 852 bp from two segments (cytochrome b and control region) of the mitochondrial genome were sequenced. In addition, orthologous sequences were obtained from eight species (11 individuals) representing other mouthbrooding lineages from Lake Tanganyika. Comparisons of sequence divergences revealed that the single Tanganyikan tribe Ectodini appears to be approximately five times older than the whole Lake Malawi cichlid species flock, suggesting that the radiation of the Tanganyikan mouthbrooding lineages took place long before the species flocks of Lakes Malawi and Victoria evolved. Seven of nine surveyed tribes of Tanganyikan cichlids appear to be approximately equally divergent, and this seems to corroborate the hypothesis of a rapid simultaneous formation of lineages at an early stage in the history of the Lake Tanganyika species flock. The close genetic relationship between the endemic Tropheus lineage and a nonendemic "Haplochromine," Astatotilapia burtoni, indicates that members of the tribe Tropheini may be the sister group of the cichlid flocks of Lakes Malawi and Victoria. The phylogenetic analyses demonstrate the monophyly of the Ectodini and identify the Cyprichromini as their sister group among the Tanganyikan cichlids. Within the tribe Ectodini the molecular data suggest both a branching pattern different than that previously proposed and a subdivision of the Ectodini into four clades, instead of the two originally described. The previously postulated model of morphological transformations believed to be responsible for the drastically different types of ecological specialization found among the Ectodini might therefore be in need of reinterpretation. Characters immediately related to foraging and nutrition seem to
Sturmbauer, C; Baric, S; Salzburger, W; Rüber, L; Verheyen, E
Water level fluctuations are important modulators of speciation processes in tropical lakes, in that they temporarily form or break down barriers to gene flow among adjacent populations and/or incipient species. Time estimates of the most recent major lowstands of the three African Great Lakes are thus crucial to infer the relative timescales of explosive speciation events in cichlid species flocks. Our approach combines geological evidence with genetic divergence data of cichlid fishes from the three Great East African Lakes derived from the fastest-evolving mtDNA segment. Thereby, we show for each of the three lakes that individuals sampled from several populations which are currently isolated by long geographic distances and/or deep water form clusters of equally closely related haplotypes. The distribution of identical or equally closely related haplotypes in a lake basin allows delineation of the extent of lake level fluctuations. Our data suggest that the same climatic phenomenon synchronized the onset of genetic divergence of lineages in all three species flocks, such that their most recent evolutionary history seems to be linked to the same external modulators of adaptive radiation. A calibration of the molecular clock of the control region was elaborated by gauging the age of the Lake Malawi species flock through the divergence among the utaka-cichlid and the mbuna-cichlid lineages to minimally 570,000 years and maximally 1 Myr. This suggests that the low-lake-level period which established the observed patterns of genetic relatedness dates back less than 57,000 years, probably even to 17,000-12,400 years ago, when Lake Victoria dried up and Lakes Malawi and Tanganyika were also low. A rapid rise of all three lakes about 11,000 years ago established the large-scale population subdivisions observed today. Over that period of time, a multitude of species originated in Lakes Malawi and Victoria with an impressive degree of morphological and ecological
Sugawara, Tohru; Terai, Yohey; Imai, Hiroo; Turner, George F; Koblmüller, Stephan; Sturmbauer, Christian; Shichida, Yoshinori; Okada, Norihiro
Many examples of the appearance of similar traits in different lineages are known during the evolution of organisms. However, the underlying genetic mechanisms have been elucidated in very few cases. Here, we provide a clear example of evolutionary parallelism, involving changes in the same genetic pathway, providing functional adaptation of RH1 pigments to deep-water habitats during the adaptive radiation of East African cichlid fishes. We determined the RH1 sequences from 233 individual cichlids. The reconstruction of cichlid RH1 pigments with 11-cis-retinal from 28 sequences showed that the absorption spectra of the pigments of nine species were shifted toward blue, tuned by two particular amino acid replacements. These blue-shifted RH1 pigments might have evolved as adaptations to the deep-water photic environment. Phylogenetic evidence indicates that one of the replacements, A292S, has evolved several times independently, inducing similar functional change. The parallel evolution of the same mutation at the same amino acid position suggests that the number of genetic changes underlying the appearance of similar traits in cichlid diversification may be fewer than previously expected.
Kudo, Yu; Nikaido, Masato; Kondo, Azusa; Suzuki, Hikoyu; Yoshida, Kohta; Kikuchi, Kiyoshi; Okada, Norihiro
Cichlid fishes in East Africa have undergone extensive adaptive radiation, which has led to spectacular diversity in their morphology and ecology. To date, genetic linkage maps have been constructed for several tilapias (riverine), Astatotilapia burtoni (Lake Tanganyika), and hybrid lines of Lake Malawi cichlids to facilitate genome-wide comparative analyses. In the present study, we constructed a genetic linkage map of the hybrid line of Lake Victoria cichlids, so that maps of cichlids from all the major areas of East Africa will be available. The genetic linkage map shown here is derived from the F2 progeny of an interspecific cross between Haplochromis chilotes and Haplochromis sauvagei and is based on 184 microsatellite and two single-nucleotide polymorphism (SNP) markers. Most of the microsatellite markers used in the present study were originally designed for other genetic linkage maps, allowing us to directly compare each linkage group (LG) among different cichlid groups. We found 25 LGs, the total length of which was 1133.2cM with an average marker spacing of about 6.09cM. Our subsequent linkage mapping analysis identified two putative sex-determining loci in cichlids. Interestingly, one of these two loci is located on cichlid LG5, on which the female heterogametic ZW locus and several quantitative trait loci (QTLs) related to adaptive evolution have been reported in Lake Malawi cichlids. We also found that V1R1 and V1R2, candidate genes for the fish pheromone receptor, are located very close to the recently detected sex-determining locus on cichlid LG5. The genetic linkage map study presented here may provide a valuable foundation for studying the chromosomal evolution of East African cichlids and the possible role of sex chromosomes in generating their genomic diversity.
Meyer, Britta S; Matschiner, Michael; Salzburger, Walter
The species-flocks of cichlid fishes in the East African Great Lakes Victoria, Malawi and Tanganyika constitute the most diverse extant adaptive radiations in vertebrates. Lake Tanganyika, the oldest of the lakes, harbors the morphologically and genetically most diverse assemblage of cichlids and contains the highest number of endemic cichlid genera of all African lakes. Based on morphological grounds, the Tanganyikan cichlid species have been grouped into 12-16 distinct lineages, so-called tribes. While the monophyly of most of the tribes is well established, the phylogenetic relationships among the tribes remain largely elusive. Here, we present a new tribal level phylogenetic hypothesis for the cichlid fishes of Lake Tanganyika that is based on the so far largest set of nuclear markers and a total alignment length of close to 18kb. Using next-generation amplicon sequencing with the 454 pyrosequencing technology, we compiled a dataset consisting of 42 nuclear loci in 45 East African cichlid species, which we subjected to maximum likelihood and Bayesian inference phylogenetic analyses. We analyzed the entire concatenated dataset and each marker individually, and performed a Bayesian concordance analysis and gene tree discordance tests. Overall, we find strong support for a position of the Oreochromini, Boulengerochromini, Bathybatini and Trematocarini outside of a clade combining the substrate spawning Lamprologini and the mouthbrooding tribes of the 'H-lineage', which are both strongly supported to be monophyletic. The Eretmodini are firmly placed within the 'H-lineage', as sister-group to the most species-rich tribe of cichlids, the Haplochromini. The phylogenetic relationships at the base of the 'H-lineage' received less support, which is likely due to high speciation rates in the early phase of the radiation. Discordance among gene trees and marker sets further suggests the occurrence of past hybridization and/or incomplete lineage sorting in the cichlid
Meyer, A; Kocher, T D; Basasibwaki, P; Wilson, A C
Lake Victoria, together with its satellite lakes, harbours roughly 200 endemic forms of cichlid fishes that are classified as 'haplochromines' and yet the lake system is less than a million years old. This 'flock' has attracted attention because of the possibility that it evolved within the lake from one ancestral species and that biologists are thus presented with a case of explosive evolution. Within the past decade, however, morphology has increasingly emphasized the view that the flock may be polyphyletic. We sequenced up to 803 base pairs of mitochondrial DNA from 14 representative Victorian species and 23 additional African species. The flock seems to be monophyletic, and is more akin to that from Lake Malawi than to species from Lake Tanganyika; in addition, it contains less genetic variation than does the human species, and there is virtually no sharing of mitochondrial DNA types among species. These results confirm that the founding event was recent.
Verheyen, E; Rüber, L; Snoeks, J; Meyer, A
The East African Lakes Tanganyika, Malawi and Victoria each harbour hundreds of endemic invertebrate and vertebrate species. Inferences about the ecological and evolutionary processes responsible for the origin of these species flocks will only be possible when they are made within historical and comparative frameworks. Specifically, the relative importance of intrinsic characteristics and extrinsic factors may offer information about the processes that drive diversification and speciation in these species. We investigated the sequence variation of a segment of the mitochondrial DNA control region of 32 populations representing all four nominal species in the three genera of eretmodine cichlids from Lake Tanganyika. Based on a phylogenetic analysis of these data we attempted to evaluate the importance of major lake level fluctuations on patterns of intralacustrine speciation. The geography of genetic variation reveals a high degree of within-lake endemism among genetically well-separated lineages distributed along the inferred shore lines of three historically intermittent lake basins. Seismic data indicate that extreme lowering of water levels in the Pleistocene caused the single Lake Tanganyika basin to split into three isolated ones. The strong phylogeographic structure of the Eretmodini, and the observation that some closely related populations occur on opposite shores of the lake, agree with this geological scenario. The three-clade-three-basin phylogeographic pattern was repeated twice within this tribe of cichlids. The phylogeographic pattern of eretmodine cichlids suggests that major fluctuations in the level of the lake have been important in shaping their adaptive radiation and speciation. The mitochondrially defined clades are in conflict with the current taxonomy of the group and suggest that there has been convergent evolution in trophic morphology, particularly in the shapes of oral teeth, taxonomically the most diagnostic characters of the three
Sefc, Kristina M
Cichlid fishes of Lake Tanganyika display a variety of mating and parental care behaviors, including polygamous and monogamous mouthbrooding and substrate breeding, cooperative breeding, as well as various alternative reproductive tactics such as sneaking and piracy. Moreover, reproductive behaviors sometimes vary within species both in space and in time. Here, I survey reports on mating and parenting behaviors of Lake Tanganyika cichlid species and address the evolution of mating and parental care patterns and sexual dimorphism. Notes on measures of sexual selection intensity and the difficulties of defining mating systems and estimating selection intensities at species level conclude the essay.
Tyers, Alexandra M.; Schiffels, Stephan; Terai, Yohey; Ngatunga, Benjamin P.; Miska, Eric A.; Durbin, Richard; Genner, Martin J.; Turner, George F.
The genomic causes and effects of divergent ecological selection during speciation are still poorly understood. Here, we report the discovery and detailed characterization of early-stage adaptive divergence of two cichlid fish ecomorphs in a small (700m diameter) isolated crater lake in Tanzania. The ecomorphs differ in depth preference, male breeding color, body shape, diet and trophic morphology. With whole genome sequences of 146 fish, we identify 98 clearly demarcated genomic ‘islands’ of high differentiation and demonstrate association of genotypes across these islands to divergent mate preferences. The islands contain candidate adaptive genes enriched for functions in sensory perception (including rhodopsin and other twilight vision associated genes), hormone signaling and morphogenesis. Our study suggests mechanisms and genomic regions that may play a role in the closely related mega-radiation of Lake Malawi. PMID:26680190
Turner, G F; Seehausen, O; Knight, M E; Allender, C J; Robinson, R L
The endemic cichlid fishes of Lakes Malawi, Tanganyika and Victoria are textbook examples of explosive speciation and adaptive radiation, and their study promises to yield important insights into these processes. Accurate estimates of species richness of lineages in these lakes, and elsewhere, will be a necessary prerequisite for a thorough comparative analysis of the intrinsic and extrinsic factors influencing rates of diversification. This review presents recent findings on the discoveries of new species and species flocks and critically appraises the relevant evidence on species richness from recent studies of polymorphism and assortative mating, generally using behavioural and molecular methods. Within the haplochromines, the most species-rich lineage, there are few reported cases of postzygotic isolation, and these are generally among allopatric taxa that are likely to have diverged a relatively long time in the past. However, many taxa, including many which occur sympatrically and do not interbreed in nature, produce viable, fertile hybrids. Prezygotic barriers are more important, and persist in laboratory conditions in which environmental factors have been controlled, indicating the primary importance of direct mate preferences. Studies to date indicate that estimates of alpha (within-site) diversity appear to be robust. Although within-species colour polymorphisms are common, these have been taken into account in previous estimates of species richness. However, overall estimates of species richness in Lakes Malawi and Victoria are heavily dependent on the assignation of species status to allopatric populations differing in male colour. Appropriate methods for testing the specific status of allopatric cichlid taxa are reviewed and preliminary results presented.
Salzburger, Walter; Niederstätter, Harald; Brandstätter, Anita; Berger, Burkhard; Parson, Walther; Snoeks, Jos; Sturmbauer, Christian
The species flocks of cichlid fishes in the East African Lakes Tanganyika, Malawi and Victoria are prime examples of adaptive radiation and explosive speciation. Several hundreds of endemic species have evolved in each of the lakes over the past several thousands to a few millions years. Sexual selection via colour-assortative mating has often been proposed as a probable causal factor for initiating and maintaining reproductive isolation. Here, we report the consequences of human-mediated admixis among differentially coloured populations of the endemic cichlid fish Tropheus moorii from several localities that have accidentally been put in sympatry in a small harbour bay in the very south of Lake Tanganyika. We analysed the phenotypes (coloration) and genotypes (mitochondrial control region and five microsatellite loci) of almost 500 individuals, sampled over 3 consecutive years. Maximum-likelihood-based parenthood analyses and Bayesian inference of population structure revealed that significantly more juveniles are the product of within-colour-morph matings than could be expected under the assumption of random mating. Our results clearly indicate a marked degree of assortative mating with respect to the different colour morphs. Therefore, we postulate that sexual selection based on social interactions and female mate choice has played an important role in the formation and maintenance of the different colour morphs in Tropheus, and is probably common in other maternally mouthbrooding cichlids as well.
Takahashi, Tetsumi; Koblmüller, Stephan
Lake Tanganyika is the oldest of the Great Ancient Lakes in the East Africa. This lake harbours about 250 species of cichlid fish, which are highly diverse in terms of morphology, behaviour, and ecology. Lake Tanganyika's cichlid diversity has evolved through explosive speciation and is treated as a textbook example of adaptive radiation, the rapid differentiation of a single ancestor into an array of species that differ in traits used to exploit their environments and resources. To elucidate the processes and mechanisms underlying the rapid speciation and adaptive radiation of Lake Tanganyika's cichlid species assemblage it is important to integrate evidence from several lines of research. Great efforts have been, are, and certainly will be taken to solve the mystery of how so many cichlid species evolved in so little time. In the present review, we summarize morphological studies that relate to the adaptive radiation of Lake Tanganyika's cichlids and highlight their importance for understanding the process of adaptive radiation. PMID:21716857
Sungani, Harold; Ngatunga, Benjamin P; Koblmüller, Stephan; Mäkinen, Tuuli; Skelton, Paul H; Genner, Martin J
It has been proposed that the fish faunas of African rivers assemble through multiple colonisation events, while lake faunas form additionally through intralacustine speciation. While this pattern has been established for many lineages, most notably cichlids, there are opportunities to further investigate the concept using phylogenies of congeneric endemic species within ancient lake catchments. The Lake Malawi catchment contains three river-spawning cyprinids of the genus Opsaridium, two of which are endemic. These species differ in body size, migratory behaviour and habitat use, but it has never previously been tested if these represent a monophyletic radiation, or have instead colonised the lake independently. We placed these species in a broader phylogeny of Opsaridium and the related genus Raiamas, including all known species from the river systems surrounding Lake Malawi. Our results suggest that each of the species has independently colonised the lake catchment, with all three taxa having well-defined sister taxa outside of the lake, and all sharing a common ancestor ∼14.9million years ago, before the Lake Malawi basin started to form ∼8.6million years ago. Additionally, the results strongly support previous observations that Opsaridium is not a monophyletic group, but instead contains Raiamas from the Congo drainage. Together these results are supportive of the concept that river fish faunas within African catchments are primarily assembled through a process of accumulation from independent origins, rather than within-catchment speciation and adaptive radiation. In light of these results we also suggest there is scope for a re-evaluation of systematics of both Opsaridium and Raiamas.
Johnson, Thomas C.; Scholz, Christopher A.; Talbot, Michael R.; Kelts, Kerry; Ricketts, R. D.; Ngobi, Gideon; Beuning, Kristina; Ssemmanda, Immacculate; McGill, J. W.
Lake Victoria is the largest lake in Africa and harbors more than 300 endemic species of haplochromine cichlid fish. Seismic reflection profiles and piston cores show that the lake not only was at a low stand but dried up completely during the Late Pleistocene, before 12,400 carbon-14 years before the present. These results imply that the rate of speciation of cichlid fish in this tropical lake has been extremely rapid.
Kuroiwa, A; Terai, Y; Kobayashi, N; Yoshida, K; Suzuki, M; Nakanishi, A; Matsuda, Y; Watanabe, M; Okada, N
Cichlid fishes in the African Great Lakes are known as a spectacular example of adaptive radiation in vertebrates. Four linkage maps have been constructed to identify the genes responsible for adaptation and speciation, and the genetic linkages of those genes are assumed to play an important role during adaptive evolution. However, it is difficult to analyze such linkages because the linkage groups of one species do not match well with those of the other species. Chromosome markers are a powerful tool for the direct identification of linkage homology between different species. We used information about the linkage map of the Lake Malawi cichlid (Labeotropheus fuelleborni/Metriaclima zebra) to isolate bacterial artificial chromosome (BAC) clones from the BAC library of Paralabidochromis chilotes, Lake Victoria. We identified 18 of 22 P. chilotes chromosomes by single- and multi-color BAC fluorescence in situ hybridization using 19 BAC clones. Comparative mapping with the chromosome markers of P. chilotes in Astatotilapia burtoni (2n = 40) from Lake Tanganyika revealed the chromosome rearrangements that have occurred in this lineage. These chromosome markers will be useful for delineating the process of genome and chromosome evolution in African species.
van Rijssel, Jacco C; Hecky, Robert E; Kishe-Machumu, Mary A; Meijer, Saskia E; Pols, Johan; van Tienderen, Kaj M; Ververs, Jan D; Wanink, Jan H; Witte, Frans
Textbook examples of adaptive radiation often show rapid morphological changes in response to environmental perturbations. East Africa's Lake Victoria, famous for its stunning adaptive radiation of cichlids, has suffered from human-induced eutrophication over the past decades. This cultural eutrophication is thought to be partly responsible for the dramatically reduced cichlid biodiversity, but climatic variability in itself might also have contributed to the eutrophication which resulted in low oxygen levels and decreased water transparency. To determine how recent environmental changes have influenced the lake and its cichlids over the past 50 years, we gathered environmental and meteorological variables and compared these with gill surface area of four cichlid species. We found that during the period of severe eutrophication and temperature increase (1980s), reduced wind speeds coincided with a reduction in oxygen levels and a decrease in both water temperature and transparency. The gill surface area in three out of the four cichlid species increased during this period which is consistent with adaptive change in response to increased hypoxia. During the 2000s, wind speeds, oxygen levels, water transparency and water temperature increased again, while cichlid gill surface area decreased. Our results imply that climatic changes and especially wind speed and direction might play a crucial role in tropical lake dynamics. The changes in Lake Victoria's water quality coincide with fluctuations in cichlid gill surface area, suggesting that these fish can respond rapidly to environmental perturbations, but also that climatic variability, together with continued eutrophication, might be detrimental to the lake's cichlid biodiversity.
Koblmüller, Stephan; Salzburger, Walter; Obermüller, Beate; Eigner, Eva; Sturmbauer, Christian; Sefc, Kristina M
The conditions of phenotypic and genetic population differentiation allow inferences about the evolution, preservation and loss of biological diversity. In Lake Tanganyika, water level fluctuations are assumed to have had a major impact on the evolution of stenotopic littoral species, though this hypothesis has not been specifically examined so far. The present study investigates whether subtly differentiated colour patterns of adjacent Tropheus moorii populations are maintained in isolation or in the face of continuous gene flow, and whether the presumed influence of water level fluctuations on lacustrine cichlids can be demonstrated in the small-scale population structure of the strictly stenotopic, littoral Tropheus. Distinct population differentiation was found even across short geographic distances and minor habitat barriers. Population splitting chronology and demographic histories comply with our expectation of old and rather stable populations on steeper sloping shore, and more recently established populations in a shallower region. Moreover, population expansions seem to coincide with lake level rises in the wake of Late Pleistocene megadroughts ~100 KYA. The imprint of hydrologic events on current population structure in the absence of ongoing gene flow suggests that phenotypic differentiation among proximate Tropheus populations evolves and persists in genetic isolation. Sporadic gene flow is effected by lake level fluctuations following climate changes and controlled by the persistence of habitat barriers during lake level changes. Since similar demographic patterns were previously reported for Lake Malawi cichlids, our data furthermore strengthen the hypothesis that major climatic events synchronized facets of cichlid evolution across the East African Great Lakes.
Mayer, W E; Tichy, H; Klein, J
The species flocks of cichlid fish in the three great East African Lakes, Victoria, Malawi, and Tanganyika, have arisen in each lake by explosive adaptive radiation. Various questions concerning their phylogeny have not yet been answered. In particular, the identity of the ancestral founder species and the monophyletic origin of the haplochromine cichlids from the East African lakes have not been established conclusively. In the present study, we used the anonymous nuclear DNA marker DXTU1 as a step towards answering these questions. A 280 bp-fragment of the DXTU1 locus was amplified by the polymerase chain reaction from East African lacustrine species, the East African riverine cichlid species Haplochromis bloyeti, H. burtoni and H. sparsidens, and other African cichlids. Sequencing revealed several indels and substitutions that were used as cladistically informative markers to support a phylogenetic tree constructed by the neighbor-joining method. The topology, although not supported by high bootstrap values, corresponds well to the geographical distribution and previous classification of the cichlids. Markers could be defined that: (i) differentiate East African from West African cichlids; (ii) distinguish the riverine and Lake Victoria/Malawi haplochromines from Lake Tanganyika cichlids; and (iii) indicate the existence of a monophyletic Lake Victoria cichlid superflock which includes haplochromines from satellite lakes and East African rivers. In order to resolve further the relationship of East African riverine and lacustrine species, mtDNA cytochrome b and control region segments were sequenced. The mtDNA-based trees support the notion of the monophyly of the Lake Victoria superflock but are ambiguous with respect to the phylogenetic position of the Lake Malawi flock.
Sültmann, H; Mayer, W E; Figueroa, F; Tichy, H; Klein, J
The recent explosive adaptive radiation of cichlids in the great lakes of Africa has attracted the attention of both morphologists and molecular biologists. To decipher the phylogenetic relationships among the various taxa within the family Cichlidae is a prerequisite for answering some fundamental questions about the nature of the speciation process. In the present study, we used the random amplification of polymorphic DNA (RAPD) technique to obtain sequence differences between selected cichlid species. We then designed specific primers based on these sequences and used them to amplify template DNA from a large number of species by the polymerase chain reaction (PCR). We sequenced the amplified products and searched the sequences for indels and shared substitutions. We identified a number of such characters at three loci--DXTU1, DXTU2, and DXTU3--and used them for phylogenetic and cladistic analysis of the relationships among the various cichlid groups. Our studies assign an outgroup position to Neotropical cichlids in relation to African cichlids, provide evidence for a sister-group relationship of tilapiines to the haplochromines, group Cyphotilapia frontosa with the lamprologines of Lake Tanganyika, place Astatoreochromis alluaudi to an outgroup position with respect to other haplochromines of Lakes Victoria and Malawi, and provide additional support for the monophyly of the remaining Lake Victoria haplochromines and the Lake Malawi haplochromines. The described approach holds great promise for further resolution of cichlid phylogeny.
McGee, Matthew D; Faircloth, Brant C; Borstein, Samuel R; Zheng, Jimmy; Darrin Hulsey, C; Wainwright, Peter C; Alfaro, Michael E
Decoupling of the upper jaw bones--jaw kinesis--is a distinctive feature of the ray-finned fishes, but it is not clear how the innovation is related to the extraordinary diversity of feeding behaviours and feeding ecology in this group. We address this issue in a lineage of ray-finned fishes that is well known for its ecological and functional diversity--African rift lake cichlids. We sequenced ultraconserved elements to generate a phylogenomic tree of the Lake Tanganyika and Lake Malawi cichlid radiations. We filmed a diverse array of over 50 cichlid species capturing live prey and quantified the extent of jaw kinesis in the premaxillary and maxillary bones. Our combination of phylogenomic and kinematic data reveals a strong association between biting modes of feeding and reduced jaw kinesis, suggesting that the contrasting demands of biting and suction feeding have strongly influenced cranial evolution in both cichlid radiations.
McGee, Matthew D.; Faircloth, Brant C.; Borstein, Samuel R.; Zheng, Jimmy; Darrin Hulsey, C.; Wainwright, Peter C.; Alfaro, Michael E.
Decoupling of the upper jaw bones—jaw kinesis—is a distinctive feature of the ray-finned fishes, but it is not clear how the innovation is related to the extraordinary diversity of feeding behaviours and feeding ecology in this group. We address this issue in a lineage of ray-finned fishes that is well known for its ecological and functional diversity—African rift lake cichlids. We sequenced ultraconserved elements to generate a phylogenomic tree of the Lake Tanganyika and Lake Malawi cichlid radiations. We filmed a diverse array of over 50 cichlid species capturing live prey and quantified the extent of jaw kinesis in the premaxillary and maxillary bones. Our combination of phylogenomic and kinematic data reveals a strong association between biting modes of feeding and reduced jaw kinesis, suggesting that the contrasting demands of biting and suction feeding have strongly influenced cranial evolution in both cichlid radiations. PMID:26763694
Wilson, A B; Noack-Kunnmann, K; Meyer, A
The growing body of empirical evidence for sympatric speciation has been complemented by recent theoretical treatments that have identified evolutionary conditions conducive to speciation in sympatry. The Neotropical Midas cichlid (Amphilophus citrinellum) fits both of the key characteristics of these models, with strong assortative mating on the basis of a colour polymorphism coupled with trophic and ecological differentiation derived from a polymorphism in their pharyngeal jaws. We used microsatellite markers and a 480 bp fragment of the mitochondrial DNA control region to study four polymorphic populations of the Midas cichlid from three crater lakes and one large lake in Nicaragua in an investigation of incipient sympatric speciation. All populations were strongly genetically differentiated on the basis of geography. We identified strong genetic separation based on colour polymorphism for populations from Lake Nicaragua and one crater lake (Lake Apoyo), but failed to find significant genetic structuring based on trophic differences and ecological niche separation in any of the four populations studied. These data support the idea that sexual selection through assortative mating contributes more strongly or earlier during speciation in sympatry than ecological separation in these cichlids. The long-term persistence of divergent cichlid ecotypes (as measured by the percentage sequence divergence between populations) in Central American crater lakes, despite a lack of fixed genetic differentiation, differs strikingly from the patterns of extremely rapid speciation in the cichlids in Africa, including its crater lakes. It is unclear whether extrinsic environmental factors or intrinsic biological differences, e.g. in the degree of phenotypic plasticity, promote different mechanisms and thereby rates of speciation of cichlid fishes from the Old and New Worlds. PMID:11413624
Abila, Romulus; Barluenga, Marta; Engelken, Johannes; Meyer, Axel; Salzburger, Walter
The approximately 500 species of the cichlid fish species flock of Lake Victoria, East Africa, have evolved in a record-setting 100,000 years and represent one of the largest adaptive radiations. We examined the population structure of the endangered cichlid species Xystichromis phytophagus from Lake Kanyaboli, a satellite lake to Lake Victoria in the Kenyan Yala wetlands. Two sets of molecular markers were analysed--sequences of the mitochondrial control region as well as six microsatellite loci--and revealed surprisingly high levels of genetic variability in this species. Mitochondrial DNA sequences failed to detect population structuring among the three sample populations. A model-based population assignment test based on microsatellite data revealed that the three populations most probably aggregate into a larger panmictic population. However, values of population pairwise FST indicated moderate levels of genetic differentiation for one population. Eleven distinct mitochondrial haplotypes were found among 205 specimens of X. phytophagus, a relatively high number compared to the total number of 54 haplotypes that were recovered from hundreds of specimens of the entire cichlid species flock of Lake Victoria. Most of the X. phytophagus mitochondrial DNA haplotypes were absent from the main Lake Victoria, corroborating the putative importance of satellite lakes as refugia for haplochromine cichlids that went extinct from the main lake in the last decades and possibly during the Late Pleistocene desiccation of Lake Victoria.
McGee, Matthew D; Borstein, Samuel R; Neches, Russell Y; Buescher, Heinz H; Seehausen, Ole; Wainwright, Peter C
Evolutionary innovations, traits that give species access to previously unoccupied niches, may promote speciation and adaptive radiation. Here, we show that such innovations can also result in competitive inferiority and extinction. We present evidence that the modified pharyngeal jaws of cichlid fishes and several marine fish lineages, a classic example of evolutionary innovation, are not universally beneficial. A large-scale analysis of dietary evolution across marine fish lineages reveals that the innovation compromises access to energy-rich predator niches. We show that this competitive inferiority shaped the adaptive radiation of cichlids in Lake Tanganyika and played a pivotal and previously unrecognized role in the mass extinction of cichlid fishes in Lake Victoria after Nile perch invasion.
Koblmüller, Stephan; Odhiambo, Elizabeth A.; Sinyinza, Danny; Sturmbauer, Christian; Sefc, Kristina M.
The largely endemic cichlid species flocks of the East African Great Lakes are among the prime examples for explosive speciation and adaptive radiation. Speciation rates differ among cichlid lineages, and the propensity to radiate has been linked to intrinsic and extrinsic factors such as sexual selection and ecological opportunity. Remarkably, only one cichlid tribe—the Boulengerochromini—comprises just a single species, Boulengerochromis microlepis, a predominantly piscivorous endemic of Lake Tanganyika and the world’s largest cichlid. While the lineage diverged from its closest relatives at the onset of the Lake Tanganyika radiation >8 MYA, mitochondrial control region sequences collected in this study dated the most recent common ancestor of B. microlepis to ~60–110 KYA. There was no evidence of phylogeographic structure in the lake-wide sample. Patterns of genetic diversity and demographic analyses were consistent with slow and steady population growth throughout the reconstructed timescale. Additionally, the shallow divergence within the species may be related to a possibly large variance in reproductive success in this highly fecund species. Trophic niche space restriction by sympatric piscivores, lack of geographic structure, low potential for sexual selection arising from the monogamous mating system and extinction may have contributed to keeping the lineage monotypic. PMID:25983338
Koblmüller, Stephan; Odhiambo, Elizabeth A; Sinyinza, Danny; Sturmbauer, Christian; Sefc, Kristina M
The largely endemic cichlid species flocks of the East African Great Lakes are among the prime examples for explosive speciation and adaptive radiation. Speciation rates differ among cichlid lineages, and the propensity to radiate has been linked to intrinsic and extrinsic factors such as sexual selection and ecological opportunity. Remarkably, only one cichlid tribe-the Boulengerochromini-comprises just a single species, Boulengerochromis microlepis, a predominantly piscivorous endemic of Lake Tanganyika and the world's largest cichlid. While the lineage diverged from its closest relatives at the onset of the Lake Tanganyika radiation >8 MYA, mitochondrial control region sequences collected in this study dated the most recent common ancestor of B. microlepis to ~60-110 KYA. There was no evidence of phylogeographic structure in the lake-wide sample. Patterns of genetic diversity and demographic analyses were consistent with slow and steady population growth throughout the reconstructed timescale. Additionally, the shallow divergence within the species may be related to a possibly large variance in reproductive success in this highly fecund species. Trophic niche space restriction by sympatric piscivores, lack of geographic structure, low potential for sexual selection arising from the monogamous mating system and extinction may have contributed to keeping the lineage monotypic.
O'Quin, Kelly E; Hofmann, Christopher M; Hofmann, Hans A; Carleton, Karen L
Phenotypic evolution may occur either through alterations to the structure of protein-coding genes or their expression. Evidence for which of these two mechanisms more commonly contribute to the evolution of a phenotype can be garnered from examples of parallel and convergent evolution. The visual system of East African cichlid fishes is an excellent system with which to address this question. Cichlid fishes from Lakes Malawi (LM) and Victoria together exhibit three diverse palettes of coexpressed opsins and several important protein-coding mutations that both shift spectral sensitivity. Here we assess both opsin expression and protein-coding diversity among cichlids from a third rift lake, Lake Tanganyika (LT). We found that Tanganyikan cichlids exhibit three palettes of coexpressed opsins that largely overlap the short-, middle-, and long-wavelength-sensitive palettes of LM cichlids. Bayesian phenotypic clustering and ancestral state reconstructions both support the parallel evolution of the short- and middle-wavelength palettes among cichlids from LT and LM. In each case, these transitions occurred from different ancestors that expressed the same long-wavelength palette. We also identified similar but distinct patterns of correlated evolution between opsin expression, diet, and lens transmittance among cichlids from LT and LM as well. In contrast to regulatory changes, we identified few functional or potentially functional mutations in the protein-coding sequences of three variable opsins, with the possible exception of the SWS1 (ultraviolet) opsin. These results underscore the important contribution that gene regulation can make to rapid phenotypic evolution and adaptation.
Miyagi, Ryutaro; Terai, Yohey
The amazing coloration shown by diverse cichlid fish not only fascinates aquarium keepers, but also receives great attention from biologists interested in speciation because of its recently-revealed role in their adaptive radiation in an African lake. We review the important role of coloration in the speciation and adaptive evolution of Lake Victoria cichlids, which have experienced adaptive radiation during a very short evolutionary period. Mature male cichlids display their colors during mate choice. The color of their skin reflects light, and the reflected light forms a color signal that is received by the visual system of females. The adaptive divergence of visual perceptions shapes and diverges colorations, to match the adapted visual perceptions. The divergence of visual perception and coloration indicates that the divergence of color signals causes reproductive isolation between species, and this process leads to speciation. Differences in color signals among coexisting species act to maintain reproductive isolation by preventing hybridization. Thus, the diversity of coloration has caused speciation and has maintained species diversity in Lake Victoria cichlids.
Ota, Kazutaka; Aibara, Mitsuto; Morita, Masaya; Awata, Satoshi; Hori, Michio; Kohda, Masanori
Alternative reproductive tactics (ARTs) are found in several Lake Tanganyika shell-brooding cichlids. Field studies were conducted in the Wonzye population to examine reproductive ecology and ARTs in the Lake Tanganyika shell-brooding cichlid Neolamprologus brevis. We discovered that this fish occurred in both rocky- and sandy-bottom habitats, but in rocky habitats, brood-caring females exclusively occurred in shell-patches that another cichlid species created. All N. brevis of both sexes in the patches were sexually mature, whereas immature males and females with unripe eggs were found frequently in sandy-bottom habitats. Males in sandy-bottom habitats were smaller, but fed more frequently and were in better somatic condition than males in the patches. Similar tendency was found in females. This indicates that N. brevis uses different habitats depending on the stage of its life history, with migration from sandy-bottom habitats to the shell-patches for reproduction. Males in the patches exhibited different behavior patterns: floating above the patches and lying in the patches. The former was larger, more aggressive, and invested less in gonads (relative to body size) than the latter. These results accord with those of other shell-brooding Lake Tanganyika cichlids with ARTs, and they therefore suggest the presence of ARTs in N. brevis. PMID:22888463
Weiss, Juliane D.; Cotterill, Fenton P. D.; Schliewen, Ulrich K.
A long history of research focused on the East Africa cichlid radiations (EAR) revealed discrepancies between mtDNA and nuclear phylogenies, suggesting that interspecific hybridisation may have been significant during the radiation of these fishes. The approximately 250 cichlid species of Lake Tanganyika have their roots in a monophyletic African cichlid assemblage, but controversies remain about the precise phylogenetic origin and placement of different lineages and consequently about L. Tanganyika colonization scenarios. 3312 AFLP loci and the mitochondrial ND2 gene were genotyped for 91 species representing almost all major lacustrine and riverine haplotilapiine east African cichlid lineages with a focus on L. Tanganyika endemics. Explicitly testing for the possibility of ancient hybridisation events, a comprehensive phylogenetic network hypothesis is proposed for the origin and diversification of L. Tanganyika cichlids. Inference of discordant phylogenetic signal strongly suggests that the genomes of two endemic L. Tanganyika tribes, Eretmodini and Tropheini, are composed of an ancient mixture of riverine and lacustrine lineages. For the first time a strong monophyly signal of all non-haplochromine mouthbrooding species endemic to L. Tanganyika (“ancient mouthbrooders”) was detected. Further, in the genomes of early diverging L. Tanganyika endemics Trematocarini, Bathybatini, Hemibatini and Boulengerochromis genetic components of other lineages belonging to the East African Radiation appear to be present. In combination with recent palaeo-geological results showing that tectonic activity in the L. Tanganyika region resulted in highly dynamic and heterogeneous landscape evolution over the Neogene and Pleistocene, the novel phylogenetic data render a single lacustrine basin as the geographical cradle of the endemic L. Tanganyika cichlid lineages unlikely. Instead a scenario of a pre-rift origin of several independent L. Tanganyika precursor lineages which
Yoshida, Kohta; Terai, Yohey; Mizoiri, Shinji; Aibara, Mitsuto; Nishihara, Hidenori; Watanabe, Masakatsu; Kuroiwa, Asato; Hirai, Hirohisa; Hirai, Yuriko; Matsuda, Yoichi; Okada, Norihiro
The endemic cichlid fishes in Lake Victoria are a model system for speciation through adaptive radiation. Although the evolution of the sex-determination system may also play a role in speciation, little is known about the sex-determination system of Lake Victoria cichlids. To understand the evolution of the sex-determination system in these fish, we performed cytogenetic analysis in 11 cichlid species from Lake Victoria. B chromosomes, which are present in addition to standard chromosomes, were found at a high prevalence rate (85%) in these cichlids. In one species, B chromosomes were female-specific. Cross-breeding using females with and without the B chromosomes demonstrated that the presence of the B chromosomes leads to a female-biased sex ratio in this species. Although B chromosomes were believed to be selfish genetic elements with little effect on phenotype and to lack protein-coding genes, the present study provides evidence that B chromosomes have a functional effect on female sex determination. FISH analysis using a BAC clone containing B chromosome DNA suggested that the B chromosomes are derived from sex chromosomes. Determination of the nucleotide sequences of this clone (104.5 kb) revealed the presence of several protein-coding genes in the B chromosome, suggesting that B chromosomes have the potential to contain functional genes. Because some sex chromosomes in amphibians and arthropods are thought to be derived from B chromosomes, the B chromosomes in Lake Victoria cichlids may represent an evolutionary transition toward the generation of sex chromosomes.
Wagner, Catherine E; McCune, Amy R; Lovette, Irby J
Lake Tanganyika, Africa's oldest lake, harbours an impressive diversity of cichlid fishes. Although diversification in its radiating groups is thought to have been initially rapid, cichlids from Lake Tanganyika show little evidence for ongoing speciation. In contrast, examples of recent divergence among sympatric colour morphs are well known in haplochromine cichlids from Lakes Malawi and Victoria. Here, we report genetic evidence for recent divergence between two sympatric Tanganyikan cichlid colour morphs. These Petrochromis morphs share mitochondrial haplotypes, yet microsatellite loci reveal that their sympatric populations form distinct genetic groups. Nuclear divergence between the two morphs is equivalent to that which arises geographically within one of the morphs over short distances and is substantially smaller than that among other sympatric species in this genus. These patterns suggest that these morphs diverged only recently, yet that barriers to gene flow exist which prevent extensive admixture despite their sympatric distribution. The morphs studied here provide an unusual example of active diversification in Lake Tanganyika's generally ancient cichlid fauna and enable comparisons of speciation processes between Lake Tanganyika and other African lakes.
Elmer, Kathryn R.; Kusche, Henrik; Lehtonen, Topi K.; Meyer, Axel
The polychromatic and trophically polymorphic Midas cichlid fish species complex (Amphilophus cf. citrinellus) is an excellent model system for studying the mechanisms of speciation and patterns of phenotypic diversification in allopatry and in sympatry. Here, we first review research to date on the species complex and the geological history of its habitat. We analyse body shape variation from all currently described species in the complex, sampled from six crater lakes (maximally 1.2–23.9 kyr old) and both great lakes in Nicaragua. We find that Midas cichlid populations in each lake have their own characteristic body shape. In lakes with multiple sympatric species of Midas cichlid, each species has a distinct body shape. Across the species complex, most body shape change relates to body depth, head, snout and mouth shape and caudal peduncle length. There is independent parallel evolution of an elongate limnetic species in at least two crater lakes. Mitochondrial genetic diversity is higher in crater lakes with multiple species. Midas cichlid species richness increases with the size and age of the crater lakes, though no such relationship exists for the other syntopic fishes. We suggest that crater lake Midas cichlids follow the predicted pattern of an adaptive radiation, with early divergence of each crater lake colonization, followed by intralacustrine diversification and speciation by ecological adaptation and sexual selection. PMID:20439280
Cichlid fish in Lake Tanganyika represent a system of adaptive radiation in which eight ancestral lineages have diversified into hundreds of species through adaptation to various niches. However, Tanganyikan cichlids have been thought to be oversaturated, that is, the species number exceeds the number of niches and ecologically equivalent and competitively even species coexist. However, recent studies have shed light on niche segregation on a finer scale among apparently equivalent species. We observed depth and substratum preferences of 15 herbivorous cichlids from four ecomorphs (i.e. grazer, browser, scraper and scooper) on a rocky littoral slope for 14 years. Depth differentiation was detected among grazers that defended feeding territories and among browsers with feeding territories. Cichlid species having no feeding territory also showed specificity on depth and substratum, resulting in habitat segregation among species that belong to the same ecomorph. Phylogenetically close species did not occupy adjacent depths, nor the opposite depth zones. Our findings suggest that apparently equivalent species of the same ecomorph coexist parapatrically along depth on a few-metre scale, or coexist with different substratum preferences on the rocky shore, and this niche segregation may have been acquired by competition between encountering equivalent species through repetitive lake-level fluctuations. PMID:28018609
Background After a volcano erupts, a lake may form in the cooled crater and become an isolated aquatic ecosystem. This makes fishes in crater lakes informative for understanding sympatric evolution and ecological diversification in barren environments. From a geological and limnological perspective, such research offers insight about the process of crater lake ecosystem establishment and speciation. In the present study we use genetic and coalescence approaches to infer the colonization history of Midas cichlid fishes (Amphilophus cf. citrinellus) that inhabit a very young crater lake in Nicaragua-the ca. 1800 year-old Lake Apoyeque. This lake holds two sympatric, endemic morphs of Midas cichlid: one with large, hypertrophied lips (~20% of the total population) and another with thin lips. Here we test the associated ecological, morphological and genetic diversification of these two morphs and their potential to represent incipient speciation. Results Gene coalescence analyses [11 microsatellite loci and mitochondrial DNA (mtDNA) sequences] suggest that crater lake Apoyeque was colonized in a single event from the large neighbouring great lake Managua only about 100 years ago. This founding in historic times is also reflected in the extremely low nuclear and mitochondrial genetic diversity in Apoyeque. We found that sympatric adult thin- and thick-lipped fishes occupy distinct ecological trophic niches. Diet, body shape, head width, pharyngeal jaw size and shape and stable isotope values all differ significantly between the two lip-morphs. The eco-morphological features pharyngeal jaw shape, body shape, stomach contents and stable isotopes (δ15N) all show a bimodal distribution of traits, which is compatible with the expectations of an initial stage of ecological speciation under disruptive selection. Genetic differentiation between the thin- and thick-lipped population is weak at mtDNA sequence (FST = 0.018) and absent at nuclear microsatellite loci (FST < 0
Until the 20th century, migrations and tribal conflicts prevented the formation of a cohesive Malawian society. Malawi is mainly comprised of a rural populace, and primary school education has only become prevalent since Malawi became independent in 1964. Shortly thereafter, Malawi adopted a new constitution and became a republic run by a president. The Malawian government is a substantially modified version of the British system and the members of the presidentially appointed cabinet are either drawn from or become members of Parliament. There are 2 judicial systems in Malawi the magisterial court headed by a 3-member Supreme Court and traditional courts created in 1964 which are based predominantly on the local court system maintained by the British prior to independence. There is only 1 authorized political party. Malawi has grown economically and has relied on expanding agricultural export as the primary vehicle for this growth. Basically, Malawi's economic outlook is favorable despite some problems which have caused a setback in the return of robust growth so apparent in the 1970s. Malawi has been basically pro-Western and has little contact with communist nations. While its political ideology is different from that of its bordering neighbors, Malawi has pursued normal relations with them and with other reighbors in its region. Relations with the US are quite good and their views on the need for economic and political stability in southern Africa are in general agreement. The 2 nations exchange dialogue and visits periodically.
Schluessel, V; Beil, O; Weber, T; Bleckmann, H
Several species have been shown to perceive symmetry as a measure of superior genetic quality, useful for assessing potential mates or mediating other visual activities such as the selection of food sources. The current study assessed whether Pseudotropheus sp. and Chiloscyllium griseum, two fish species from distantly related groups, possess symmetry perception. In alternative two choice experiments, individuals were tested for spontaneous preferences and trained to discriminate between abstract symmetrical and asymmetrical stimulus pairs. Pair discriminations were followed by extensive categorization experiments. Transfer tests elucidated whether bilaterally symmetrical and rotationally symmetrical stimuli could be distinguished. Sharks were also tested for the degree of dissimilarity between two symbols that could still be detected. While sharks showed both a spontaneous preference for symmetry as well as remarkable discrimination abilities by succeeding in all of the presented tasks, cichlids showed no spontaneous preference, had difficulties in discriminating between symbols and performed poorly in the categorization experiments. Sharks distinguished between bilaterally and rotationally symmetrical stimuli and easily differentiated between a four-armed cross (all arms 90° apart) and a cross where one of the arms was only 45° spaced from the one next to it. Performance did not decline when the separation was extended to 70°, but was significantly reduced at an 80° separation. Results indicate that the ability for symmetry perception varies across fish species and individuals, whereby some can detect even subtle differences in this respect.
Hulsey, C.D; Mims, M.C; Streelman, J.T
Constraints on form should determine how organisms diversify. Owing to competition for the limited space within the body, investment in adjacent structures may frequently represent an evolutionary compromise. For example, evolutionary trade-offs between eye size and jaw muscles in cichlid fish of the African great lakes are thought to represent a constructional constraint that influenced the diversification of these assemblages. To test the evolutionary independence of these structures in Lake Malawi cichlid fish, we measured the mass of the three major adductor mandibulae (AM) muscles and determined the eye volume in 41 species. Using both traditional and novel methodologies to control for resolved and unresolved phylogenetic relationships, we tested the evolutionary independence of these four structures. We found that evolutionary change in the AM muscles was positively correlated, suggesting that competition for space in the head has not influenced diversification among these jaw muscles. Furthermore, there was no negative relationship between change in total AM muscle mass and eye volume, indicating that there has been little effect of the evolution of eye size on AM evolution in Lake Malawi cichlids. The comparative approach used here should provide a robust method to test whether constructional constraints frequently limit phenotypic change in adaptive radiations. PMID:17519189
Elmer, Kathryn R; Reggio, Chiara; Wirth, Thierry; Verheyen, Erik; Salzburger, Walter; Meyer, Axel
The Great Lakes region of East Africa, including Lake Victoria, is the center of diversity of the mega-diverse cichlid fishes (Perciformes: Teleostei). Paleolimnological evidence indicates dramatic desiccation of this lake ca. 18,000-15,000 years ago. Consequently, the hundreds of extant endemic haplochromine species in the lake must have either evolved since then or refugia must have existed, within that lake basin or elsewhere, from which Lake Victoria was recolonized. We studied the population history of the Lake Victoria region superflock (LVRS) of haplochromine cichlids based on nuclear genetic analysis (12 microsatellite loci from 400 haplochomines) of populations from Lake Kivu, Lake Victoria, and the connected and surrounding rivers and lakes. Population genetic analyses confirmed that Lake Kivu haplochromines colonized Lake Victoria. Coalescent analyses show a 30- to 50-fold decline in the haplochromine populations of Lake Victoria, Lake Kivu, and the region ca. 18,000-15,000 years ago. We suggest that this coincides with drastic climatic and geological changes in the late Pleistocene. The most recent common ancestor of the Lake Victoria region haplochromines was estimated to have existed about 4.5 million years ago, which corresponds to the first radiation of cichlids in Lake Tanganyika and the origin of the tribe Haplochrominii. This relatively old evolutionary origin may explain the high levels of polymorphism still found in modern haplochromines. This degree of polymorphism might have acted as a "genetic reservoir" that permitted the explosive radiation of hundreds of haplochromines and their array of contemporary adaptive morphologies.
Rüber, Lukas; Verheyen, Erik; Meyer, Axel
The current phylogenetic hypothesis for the endemic Lake Tanganyika cichlid fishes of the tribe Eretmodini is based solely on morphology and suggests that more complex trophic morphologies derived only once from a less specialized ancestral condition. A molecular phylogeny of eretmodine cichlids based on partial mitochondrial DNA cytochrome b and control-region sequences was used to reconstruct the evolutionary sequence of trophic adaptations and to test alternative models of morphological divergence. The six mitochondrial lineages found disagree with the current taxonomy and the morphology-based phylogeny. Mitochondrial lineages with similar trophic morphologies are not grouped monophyletically but are typically more closely related to lineages with different trophic phenotypes currently assigned to other genera. Our results indicate multiple independent origins of similar trophic specializations in these cichlids. A pattern of repeated divergent morphological evolution becomes apparent when the phylogeography of the mitochondrial haplotypes is analyzed in the context of the geological and paleoclimatological history of Lake Tanganyika. In more than one instance within Lake Tanganyika, similar morphological divergence of dentitional traits occurred in sympatric species pairs. Possibly, resource-based divergent selective regimes led to resource partitioning and brought about similar trophic morphologies independently and repeatedly. PMID:10468591
Meyer, Britta S; Indermaur, Adrian; Ehrensperger, Xenia; Egger, Bernd; Banyankimbona, Gaspard; Snoeks, Jos; Salzburger, Walter
The species flocks of cichlid fishes in the East African Great Lakes are the largest vertebrate adaptive radiations in the world and illustrious textbook examples of convergent evolution between independent species assemblages. Although recent studies suggest some degrees of genetic exchange between riverine taxa and the lake faunas, not a single cichlid species is known from Lakes Tanganyika, Malawi and Victoria that is derived from the radiation associated with another of these lakes. Here, we report the discovery of a haplochromine cichlid species in Lake Tanganyika, which belongs genetically to the species flock of haplochromines of the Lake Victoria region. The new species colonized Lake Tanganyika only recently, suggesting that faunal exchange across watersheds and, hence, between isolated ichthyofaunas, is more common than previously thought.
Kassam, Daud; Seki, Shingo; Horic, Michio; Yamaoka, Kosaku
The apparent inter-lake morphological similarity among East African Great Lakes' cichlid species/genera has left evolutionary biologists asking whether such similarity is due to sharing of common ancestor or mere convergent evolution. In order to answer such question, we first used Geometric Morphometrics, GM, to quantify morphological similarity and then subsequently used Amplified Fragment Length Polymorphism, AFLP, to determine if similar morphologies imply shared ancestry or convergent evolution. GM revealed that not all presumed morphological similar pairs were indeed similar, and the dendrogram generated from AFLP data indicated distinct clusters corresponding to each lake and not inter-lake morphological similar pairs. Such results imply that the morphological similarity is due to convergent evolution and not shared ancestry. The congruency of GM and AFLP generated dendrograms imply that GM is capable of picking up phylogenetic signal, and thus GM can be potential tool in phylogenetic systematics.
Vanhove, Maarten P M; Pariselle, Antoine; Van Steenberge, Maarten; Raeymaekers, Joost A M; Hablützel, Pascal I; Gillardin, Céline; Hellemans, Bart; Breman, Floris C; Koblmüller, Stephan; Sturmbauer, Christian; Snoeks, Jos; Volckaert, Filip A M; Huyse, Tine
The stunning diversity of cichlid fishes has greatly enhanced our understanding of speciation and radiation. Little is known about the evolution of cichlid parasites. Parasites are abundant components of biodiversity, whose diversity typically exceeds that of their hosts. In the first comprehensive phylogenetic parasitological analysis of a vertebrate radiation, we study monogenean parasites infecting tropheine cichlids from Lake Tanganyika. Monogeneans are flatworms usually infecting the body surface and gills of fishes. In contrast to many other parasites, they depend only on a single host species to complete their lifecycle. Our spatially comprehensive combined nuclear-mitochondrial DNA dataset of the parasites covering almost all tropheine host species (N = 18), reveals species-rich parasite assemblages and shows consistent host-specificity. Statistical comparisons of host and parasite phylogenies based on distance and topology-based tests demonstrate significant congruence and suggest that host-switching is rare. Molecular rate evaluation indicates that species of Cichlidogyrus probably diverged synchronically with the initial radiation of the tropheines. They further diversified through within-host speciation into an overlooked species radiation. The unique life history and specialisation of certain parasite groups has profound evolutionary consequences. Hence, evolutionary parasitology adds a new dimension to the study of biodiversity hotspots like Lake Tanganyika.
Vanhove, Maarten P. M.; Pariselle, Antoine; Van Steenberge, Maarten; Raeymaekers, Joost A. M.; Hablützel, Pascal I.; Gillardin, Céline; Hellemans, Bart; Breman, Floris C.; Koblmüller, Stephan; Sturmbauer, Christian; Snoeks, Jos; Volckaert, Filip A. M.; Huyse, Tine
The stunning diversity of cichlid fishes has greatly enhanced our understanding of speciation and radiation. Little is known about the evolution of cichlid parasites. Parasites are abundant components of biodiversity, whose diversity typically exceeds that of their hosts. In the first comprehensive phylogenetic parasitological analysis of a vertebrate radiation, we study monogenean parasites infecting tropheine cichlids from Lake Tanganyika. Monogeneans are flatworms usually infecting the body surface and gills of fishes. In contrast to many other parasites, they depend only on a single host species to complete their lifecycle. Our spatially comprehensive combined nuclear-mitochondrial DNA dataset of the parasites covering almost all tropheine host species (N = 18), reveals species-rich parasite assemblages and shows consistent host-specificity. Statistical comparisons of host and parasite phylogenies based on distance and topology-based tests demonstrate significant congruence and suggest that host-switching is rare. Molecular rate evaluation indicates that species of Cichlidogyrus probably diverged synchronically with the initial radiation of the tropheines. They further diversified through within-host speciation into an overlooked species radiation. The unique life history and specialisation of certain parasite groups has profound evolutionary consequences. Hence, evolutionary parasitology adds a new dimension to the study of biodiversity hotspots like Lake Tanganyika. PMID:26335652
Malinsky, Milan; Keller, Irene; Fan, Shaohua; Simakov, Oleg; Ng, Alvin Y.; Lim, Zhi Wei; Bezault, Etienne; Turner-Maier, Jason; Johnson, Jeremy; Alcazar, Rosa; Noh, Hyun Ji; Russell, Pamela; Aken, Bronwen; Alföldi, Jessica; Amemiya, Chris; Azzouzi, Naoual; Baroiller, Jean-François; Barloy-Hubler, Frederique; Berlin, Aaron; Bloomquist, Ryan; Carleton, Karen L.; Conte, Matthew A.; D'Cotta, Helena; Eshel, Orly; Gaffney, Leslie; Galibert, Francis; Gante, Hugo F.; Gnerre, Sante; Greuter, Lucie; Guyon, Richard; Haddad, Natalie S.; Haerty, Wilfried; Harris, Rayna M.; Hofmann, Hans A.; Hourlier, Thibaut; Hulata, Gideon; Jaffe, David B.; Lara, Marcia; Lee, Alison P.; MacCallum, Iain; Mwaiko, Salome; Nikaido, Masato; Nishihara, Hidenori; Ozouf-Costaz, Catherine; Penman, David J.; Przybylski, Dariusz; Rakotomanga, Michaelle; Renn, Suzy C. P.; Ribeiro, Filipe J.; Ron, Micha; Salzburger, Walter; Sanchez-Pulido, Luis; Santos, M. Emilia; Searle, Steve; Sharpe, Ted; Swofford, Ross; Tan, Frederick J.; Williams, Louise; Young, Sarah; Yin, Shuangye; Okada, Norihiro; Kocher, Thomas D.; Miska, Eric A.; Lander, Eric S.; Venkatesh, Byrappa; Fernald, Russell D.; Meyer, Axel; Ponting, Chris P.; Streelman, J. Todd; Lindblad-Toh, Kerstin; Seehausen, Ole; Di Palma, Federica
Cichlid fishes are famous for large, diverse and replicated adaptive radiations in the Great Lakes of East Africa. To understand the molecular mechanisms underlying cichlid phenotypic diversity, we sequenced the genomes and transcriptomes of five lineages of African cichlids: the Nile tilapia (Oreochromis niloticus), an ancestral lineage with low diversity; and four members of the East African lineage: Neolamprologus brichardi/pulcher (older radiation, Lake Tanganyika), Metriaclima zebra (recent radiation, Lake Malawi), Pundamilia nyererei (very recent radiation, Lake Victoria), and Astatotilapia burtoni (riverine species around Lake Tanganyika). We found an excess of gene duplications in the East African lineage compared to tilapia and other teleosts, an abundance of non-coding element divergence, accelerated coding sequence evolution, expression divergence associated with transposable element insertions, and regulation by novel microRNAs. In addition, we analysed sequence data from sixty individuals representing six closely related species from Lake Victoria, and show genome-wide diversifying selection on coding and regulatory variants, some of which were recruited from ancient polymorphisms. We conclude that a number of molecular mechanisms shaped East African cichlid genomes, and that amassing of standing variation during periods of relaxed purifying selection may have been important in facilitating subsequent evolutionary diversification. PMID:25186727
Background Phenotypic integration among different anatomical parts of the head is a common phenomenon across vertebrates. Interestingly, despite centuries of research into the factors that contribute to the existing variation in brain size among vertebrates, little is known about the role of phenotypic integration in brain size diversification. Here we used geometric morphometrics on the morphologically diverse Tanganyikan cichlids to investigate phenotypic integration across key morphological aspects of the head. Then, while taking the effect of shared ancestry into account, we tested if head shape was associated with brain size while controlling for the potentially confounding effect of feeding strategy. Results The shapes of the anterior and posterior parts of the head were strongly correlated, indicating that the head represents an integrated morphological unit in Lake Tanganyika cichlids. After controlling for phylogenetic non-independence, we also found evolutionary associations between head shape, brain size and feeding ecology. Conclusions Geometric morphometrics and phylogenetic comparative analyses revealed that the anterior and posterior parts of the head are integrated, and that head morphology is associated with brain size and feeding ecology in Tanganyikan cichlid fishes. In light of previous results on mammals, our results suggest that the influence of phenotypic integration on brain diversification is a general process. PMID:24593160
Van Bocxlaer, Bert; Hunt, Gene
Evolutionary processes leading to adaptive radiation regularly occur too fast to be accurately recorded in the fossil record but too slowly to be readily observed in living biota. The study of evolutionary radiations is thereby confronted with an epistemological gap between the timescales and approaches used by neontologists and paleontologists. Here we report on an ongoing radiation of extant Bellamya species (n = 4) from the African Rift Lake Malawi that provides an unusual opportunity to bridge this gap. The substantial molecular differentiation in this monophyletic Bellamya clade has arisen since Late Pleistocene megadroughts in the Malawi Basin caused by climate change. Morphological time-series analysis of a high-resolution, radiocarbon-dated sequence of 22 faunas spanning the Holocene documents stasis up to the middle Holocene in all traits studied (shell height, number of whorls, and two variables obtained from geometric morphometrics). Between deposition of the last fossil fauna (∼5 ka) and the present day, a drastic increase in morphological disparity was observed (3.7–5.8 times) associated with an increase in species diversity. Comparison of the rates of morphological evolution obtained from the paleontological time-series with phylogenetic rates indicates that the divergence in two traits could be reconstructed with the slow rates documented in the fossils, that one trait required a rate reduction (stabilizing selection), and the other faster rates (divergent selection). The combined paleontological and comparative approach taken here allows recognition that morphological stasis can be the dominant evolutionary pattern within species lineages, even in very young and radiating clades. PMID:23924610
Theis, Anya; Ronco, Fabrizia; Indermaur, Adrian; Salzburger, Walter; Egger, Bernd
Divergent natural selection acting in different habitats may build up barriers to gene flow and initiate speciation. This speciation continuum can range from weak or no divergence to strong genetic differentiation between populations. Here, we focus on the early phases of adaptive divergence in the East African cichlid fish Astatotilapia burtoni, which occurs in both Lake Tanganyika (LT) and inflowing rivers. We first assessed the population structure and morphological differences in A. burtoni from southern LT. We then focused on four lake-stream systems and quantified body shape, ecologically relevant traits (gill raker and lower pharyngeal jaw) as well as stomach contents. Our study revealed the presence of several divergent lake-stream populations that rest at different stages of the speciation continuum, but show the same morphological and ecological trajectories along the lake-stream gradient. Lake fish have higher bodies, a more superior mouth position, longer gill rakers and more slender pharyngeal jaws, and they show a plant/algae and zooplankton-biased diet, whereas stream fish feed more on snails, insects and plant seeds. A test for reproductive isolation between closely related lake and stream populations did not detect population-assortative mating. Analyses of F1 offspring reared under common garden conditions indicate that the detected differences in body shape and gill raker length do not constitute pure plastic responses to different environmental conditions, but also have a genetic basis. Taken together, the A. burtoni lake-stream system constitutes a new model to study the factors that enhance and constrain progress towards speciation in cichlid fishes.
Background Elucidation of the mechanisms driving speciation requires detailed knowledge about the phylogenetic relationships and phylogeography of the incipient species within their entire ranges as well as their colonization history. The Midas cichlid species complex Amphilophus spp. has been proven to be a powerful model system for the study of ecological specialization, sexual selection and the mechanisms of sympatric speciation. Here we present a comprehensive and integrative phylogeographic analysis of the complete Midas Cichlid species complex in Nicaragua (> 2000 individuals) covering the entire distributional range, using two types of molecular markers (the mitochondrial DNA control region and 15 microsatellites). We investigated the majority of known lake populations of this species complex and reconstructed their colonization history in order to distinguish between alternative speciation scenarios. Results We found that the large lakes contain older and more diverse Midas Cichlid populations, while all crater lakes hold younger and genetically less variable species assemblages. The large lakes appear to have repeatedly acted as source populations for all crater lakes, and our data indicate that faunal exchange among crater lakes is extremely unlikely. Despite their very recent (often only a few thousand years old) and common origin from the two large Nicaraguan lakes, all crater lake Midas Cichlid radiations underwent independent, but parallel, evolution, and comprise distinct genetic units. Indeed several of these crater lakes contain multiple genetically distinct incipient species that most likely arose through sympatric speciation. Several crater lake radiations can be traced back to a single ancestral line, but some appear to have more than one founding lineage. The timing of the colonization(s) of each crater lake differs, although most of them occurred more (probably much more) recently than 20,000 years ago. Conclusion The genetic differentiation
Takeda, Miyuki; Kusumi, Junko; Mizoiri, Shinji; Aibara, Mitsuto; Mzighani, Semvua Isa; Sato, Tetsu; Terai, Yohey; Okada, Norihiro; Tachida, Hidenori
The approximately 700 species of cichlids found in Lake Victoria in East Africa are thought to have evolved over a short period of time, and they represent one of the largest known examples of adaptive radiation. To understand the processes that are driving this spectacular radiation, we must determine the present genetic structure of these species and elucidate how this structure relates to the ecological conditions that caused their adaptation. We analyzed the genetic structure of two pelagic and seven littoral species sampled from the southeast area of Lake Victoria using sequences from the mtDNA control region and 12 microsatellite loci as markers. Using a Bayesian model-based clustering method to analyze the microsatellite data, we separated these nine species into four groups: one group composed of pelagic species and another three groups composed mainly of rocky-shore species. Furthermore, we found significant levels of genetic variation between species within each group at both marker loci using analysis of molecular variance (AMOVA), although the nine species often shared mtDNA haplotypes. We also found significant levels of genetic variation between populations within species. These results suggest that initial groupings, some of which appear to have been related to habitat differences, as well as divergence between species within groups took place among the cichlid species of Lake Victoria.
Van Damme, D.; Gautier, A.
In terminal Pliocene-early Pleistocene times, part of the Malawi Basin was occupied by paleo-lake Chiwondo. Molluscan biostratigraphy situates this freshwater lake either in the East African wet phase between 2.7-2.4 Ma or that of 2.0-1.8 Ma. In-lake divergent evolution remained restricted to a few molluscan taxa and was very modest. The lacustrine Chiwondo fauna went extinct at the beginning of the Pleistocene. The modern Lake Malawi malacofauna is depauperate and descends from ubiquistic southeast African taxa and some Malawi basin endemics that invaded the present lake after the Late Pleistocene mega-droughts. The Pleistocene aridity crises caused dramatic changes, affecting the malacofauna of all East African lakes. All lacustrine endemic faunas that had evolved in the Pliocene rift lakes, such as paleo-lake Chiwondo, became extinct. In Lake Tanganyika, the freshwater ecosystem did not crash as in other lakes, but the environmental changes were sufficiently important to trigger a vast radiation. All African endemic lacustrine molluscan clades that are the result of in-lake divergence are hence geologically young, including the vast Lavigeria clade in Lake Tanganyika (ca. 43 species).
Danley, Patrick D.; Husemann, Martin; Ding, Baoqing; DiPietro, Lyndsay M.; Beverly, Emily J.; Peppe, Daniel J.
The cichlid fishes of the East African Great Lakes are the largest extant vertebrate radiation identified to date. These lakes and their surrounding waters support over 2,000 species of cichlid fish, many of which are descended from a single common ancestor within the past 10 Ma. The extraordinary East African cichlid diversity is intricately linked to the highly variable geologic and paleoclimatic history of this region. Greater than 10 Ma, the western arm of the East African rift system began to separate, thereby creating a series of rift basins that would come to contain several water bodies, including the extremely deep Lakes Tanganyika and Malawi. Uplifting associated with this rifting backponded many rivers and created the extremely large, but shallow Lake Victoria. Since their creation, the size, shape, and existence of these lakes have changed dramatically which has, in turn, significantly influenced the evolutionary history of the lakes' cichlids. This paper reviews the geologic history and paleoclimate of the East African Great Lakes and the impact of these forces on the region's endemic cichlid flocks. PMID:22888465
Danley, Patrick D; Husemann, Martin; Ding, Baoqing; Dipietro, Lyndsay M; Beverly, Emily J; Peppe, Daniel J
The cichlid fishes of the East African Great Lakes are the largest extant vertebrate radiation identified to date. These lakes and their surrounding waters support over 2,000 species of cichlid fish, many of which are descended from a single common ancestor within the past 10 Ma. The extraordinary East African cichlid diversity is intricately linked to the highly variable geologic and paleoclimatic history of this region. Greater than 10 Ma, the western arm of the East African rift system began to separate, thereby creating a series of rift basins that would come to contain several water bodies, including the extremely deep Lakes Tanganyika and Malawi. Uplifting associated with this rifting backponded many rivers and created the extremely large, but shallow Lake Victoria. Since their creation, the size, shape, and existence of these lakes have changed dramatically which has, in turn, significantly influenced the evolutionary history of the lakes' cichlids. This paper reviews the geologic history and paleoclimate of the East African Great Lakes and the impact of these forces on the region's endemic cichlid flocks.
Kusche, Henrik; Recknagel, Hans; Elmer, Kathryn Rebecca; Meyer, Axel
A common pattern of adaptive diversification in freshwater fishes is the repeated evolution of elongated open water (limnetic) species and high-bodied shore (benthic) species from generalist ancestors. Studies on phenotype-diet correlations have suggested that population-wide individual specialization occurs at an early evolutionary and ecological stage of divergence and niche partitioning. This variable restricted niche use across individuals can provide the raw material for earliest stages of sympatric divergence. We investigated variation in morphology and diet as well as their correlations along the benthic-limnetic axis in an extremely young Midas cichlid species, Amphilophus tolteca, endemic to the Nicaraguan crater lake Asososca Managua. We found that A. tolteca varied continuously in ecologically relevant traits such as body shape and lower pharyngeal jaw morphology. The correlation of these phenotypes with niche suggested that individuals are specialized along the benthic-limnetic axis. No genetic differentiation within the crater lake was detected based on genotypes from 13 microsatellite loci. Overall, we found that individual specialization in this young crater lake species encompasses the limnetic-as well as the benthic macro-habitat. Yet there is no evidence for any diversification within the species, making this a candidate system for studying what might be the early stages preceding sympatric divergence. A common pattern of adaptive diversification in freshwater fishes is the repeated evolution of open water (limnetic) species and of shore (benthic) species. Individual specialization can reflect earliest stages of evolutionary and ecological divergence. We here demonstrate individual specialization along the benthic–limnetic axis in a young adaptive radiation of crater lake cichlid fishes. PMID:24772288
Ellis, Geoffrey S.; Barry J. Katz,; Christopher A. Scholz,; Peter K. Swart,
This study examines the relationship between depositional environment and sedimentary organic geochemistry in Lake Malawi, East Africa, and evaluates the relative significance of the various processes that control sedimentary organic matter (OM) in lacustrine systems. Total organic carbon (TOC) concentrations in recent sediments from Lake Malawi range from 0.01 to 8.80 wt% and average 2.83 wt% for surface sediments and 2.35 wt% for shallow core sediments. Hydrogen index (HI) values as determined by Rock-Eval pyrolysis range from 0 to 756 mg HC g−1 TOC and average 205 mg HC g−1 TOC for surface sediments and 228 mg HC g−1 TOC for shallow core samples. On average, variations in primary productivity throughout the lake may account for ~33% of the TOC content in Lake Malawi sediments (as much as 1 wt% TOC), and have little or no impact on sedimentary HI values. Similarly, ~33% to 66% of the variation in TOC content in Lake Malawi sediments appears to be controlled by anoxic preservation of OM (~1–2 wt% TOC), although some component of the water depth–TOC relationship may be due to physical sediment transport processes. Furthermore, anoxic preservation has a minimal effect on HI values in Lake Malawi sediments. Dilution of OM by inorganic sediment may account for ~16% of variability in TOC content in Lake Malawi sediments (~0.5 wt% TOC). The effect of inputs of terrestrial sediment on the organic character of surface sediments in these lakes is highly variable, and appears to be more closely related to the local depositional environment than the regional flux of terrestrial OM. Total nitrogen and TOC content in surface sediments collected throughout the lake are found to be highly correlated (r2 = 0.95), indicating a well-homogenized source of OM to the lake bottom. The recurring suspension and deposition of terrestrial sediment may account for significant amounts of OM deposited in offshore regions of the lake. This process effectively separates denser
Schultheiß, Roland; Van Bocxlaer, Bert; Wilke, Thomas; Albrecht, Christian
Studies on environmental changes provide important insights into modes of speciation, into the (adaptive) reoccupation of ecological niches and into species turnover. Against this background, we here examine the history of the gastropod genus Lanistes in the African Rift Lake Malawi, guided by four general evolutionary scenarios, and compare it with patterns reported from other endemic Malawian rift taxa. Based on an integrated approach using a mitochondrial DNA phylogeny and a trait-specific molecular clock in combination with insights from the fossil record and palaeoenvironmental data, we demonstrate that the accumulation of extant molecular diversity in the endemic group did not start before approximately 600 000 years ago from a single lineage. Fossils of the genus from the Malawi Rift, however, are over one million years older. We argue that severe drops in the lake level of Lake Malawi in the Pleistocene offer a potential explanation for this pattern. Our results also challenge previously established phylogenetic relationships within the genus by revealing parallel evolution and providing evidence that the endemic Lanistes species are not restricted to the lake proper but are present throughout the Malawi Rift. PMID:19439440
Takahashi, T; Sota, T; Hori, M
Phenotypic polymorphisms can be applied to study the micro-evolutionary forces that maintain genetic variation and can mediate speciation, but it can be difficult to determine the genetic basis of polymorphisms. Recently, restriction-site-associated DNA (RAD) sequencing has become popular, which can easily produce multiple single nucleotide polymorphisms from whole genomes. Here, we combined RAD sequencing, allele-specific PCR and Sanger sequencing to determine the genetic basis underlying male colour dimorphism of a Lake Tanganyika cichlid fish, Cyprichromis leptosoma. Our analyses using both a cross-family (two parents and 12 F2 males) and 64 wild individuals do not contradict a hypothesis that two alleles of one-locus control male colour dimorphism. Also, the locus may be located on a genome region that experiences reduced levels of recombination. Although more analyses will be needed to conclude these findings, this study is the first to suggest the genetic basis of a colour polymorphism using RAD sequencing.
Muschick, Moritz; Nosil, Patrik; Roesti, Marius; Dittmann, Marie Theres; Harmon, Luke; Salzburger, Walter
Adaptive radiation (AR) is a key process in the origin of organismal diversity. However, the evolution of trait disparity in connection with ecological specialization is still poorly understood. Available models for vertebrate ARs predict that diversification occurs in the form of temporal stages driven by different selective forces. Here, we investigate the AR of cichlid fishes in East African Lake Tanganyika and use macroevolutionary model fitting to evaluate whether diversification happened in temporal stages. Six trait complexes, for which we also provide evidence of their adaptiveness, are analysed with comparative methods: body shape, pharyngeal jaw shape, gill raker traits, gut length, brain weight and body coloration. Overall, we do not find strong evidence for the ‘stages model’ of AR. However, our results suggest that trophic traits diversify earlier than traits implicated in macrohabitat adaptation and that sexual communication traits (i.e. coloration) diversify late in the radiation. PMID:25274371
Fruciano, Carmelo; Franchini, Paolo; Kovacova, Viera; Elmer, Kathryn R.; Henning, Frederico; Meyer, Axel
Our understanding of how biological diversity arises is limited, especially in the case of speciation in the face of gene flow. Here we investigate the genomic basis of adaptive traits, focusing on a sympatrically diverging species pair of crater lake cichlid fishes. We identify the main quantitative trait loci (QTL) for two eco-morphological traits: body shape and pharyngeal jaw morphology. These traits diverge in parallel between benthic and limnetic species in the repeated adaptive radiations of this and other fish lineages. Remarkably, a single chromosomal region contains the highest effect size QTL for both traits. Transcriptomic data show that the QTL regions contain genes putatively under selection. Independent population genomic data corroborate QTL regions as areas of high differentiation between the sympatric sister species. Our results provide empirical support for current theoretical models that emphasize the importance of genetic linkage and pleiotropy in facilitating rapid divergence in sympatry. PMID:27597183
Bezault, Etienne; Mwaiko, Salome; Seehausen, Ole
Adaptive radiation is usually thought to be associated with speciation, but the evolution of intraspecific polymorphisms without speciation is also possible. The radiation of cichlid fish in Lake Victoria (LV) is perhaps the most impressive example of a recent rapid adaptive radiation, with 600+ very young species. Key questions about its origin remain poorly characterized, such as the importance of speciation versus polymorphism, whether species persist on evolutionary time scales, and if speciation happens more commonly in small isolated or in large connected populations. We used 320 individuals from 105 putative species from Lakes Victoria, Edward, Kivu, Albert, Nabugabo and Saka, in a radiation-wide amplified fragment length polymorphism (AFLP) genome scan to address some of these questions. We demonstrate pervasive signatures of speciation supporting the classical model of adaptive radiation associated with speciation. A positive relationship between the age of lakes and the average genomic differentiation of their species, and a significant fraction of molecular variance explained by above-species level taxonomy suggest the persistence of species on evolutionary time scales, with radiation through sequential speciation rather than a single starburst. Finally the large gene diversity retained from colonization to individual species in every radiation suggests large effective population sizes and makes speciation in small geographical isolates unlikely.
Booton, G C; Kaufman, L; Chandler, M; Oguto-Ohwayo, R; Duan, W; Fuerst, P A
The nucleotide sequences of the first internal transcribed spacer (ITS-1) of the ribosomal RNA gene cluster have been determined for 11 species of closely related endemic cichlid fishes of the Lake Victoria region (LVR) and 6 related East African cichlids. The ITS-1 sequences confirmed independently derived basal phylogenies, but provide limited insight within this species flock. The line leading to Pseudocrenilabrus multicolor arose early, close to the divergence event that separated the tilapiine and haplochromine tribes of the "African Group" of the family Cichlidae. In this phylogeny, Astatoreochromis alluaudi and the riverine Astatotilapia burtoni are sister taxa, which together are a sister group to a monophyletic assemblage including both Lake Victoria and Lake Edward taxa. The ITS-1 data support the monophyly of haplochromine genera across lakes. Since Lake Victoria is believed to have been dry between 14, 500 and 12,400 BPE, the modern assemblage must have been derived from reinvasion by the products of earlier cladogenesis events. Thus, although the regional superflock is monophyletic, the haplochromines of Lake Victoria itself did not evolve in situ from a single ancestor.
Sturmbauer, Christian; Salzburger, Walter; Duftner, Nina; Schelly, Robert; Koblmüller, Stephan
Lake Tanganyika comprises a cichlid species flock with substrate-breeding and mouthbrooding lineages. While sexual selection via mate choice on male mating color is thought to boost speciation rates in mouthbrooding cichlids, this is not the case in substrate-breeding lamprologines, which mostly form stable pairs and lack sexual dichromatism. We present a comprehensive reconstruction of the evolution of the cichlid tribe Lamprologini, based upon mtDNA sequences and multilocus nuclear DNA (AFLP) markers. Twelve mtDNA clades were identified, seven of which were corroborated by the AFLP tree. The radiation is likely to have started about 5.3 MYA, contemporarily with that of the mouthbrooding C-lineage, and probably triggered by the onset of deep-water conditions in Lake Tanganyika. Neither the Congo- nor the Malagarazi River species form the most ancestral branch. Several conflicts in the mtDNA phylogeny with taxonomic assignments based upon color, eco-morphology and behavior could be resolved and complemented by the AFLP analysis. Introgressive hybridization upon secondary contact seems to be the most likely cause for paraphyly of taxa due to mtDNA capture in species involving brood-care helpers, while accidental hybridization best explains the para- or polyphyly of several gastropod shell breeders. Taxonomic error or paraphyly due to the survival of ancestral lineages appear responsible for inconsistencies in the genera Lamprologus and Neolamprologus. PMID:20601006
de Zeeuw, Marnix P.; Westbroek, Irene; van Oijen, Martien J.P.; Witte, Frans
Abstract Two new species of zooplanktivorous haplochromine cichlids from Lake Victoria, Tanzania, are described and illustrated. These species closely resemble each other. Their affinities to other zooplanktivorous haplochromines from Lake Victoria are discussed. Haplochromis argens sp. n., which featured under nicknames (mainly Haplochromis “argens”) in more than 50 papers, was caught both in the Mwanza Gulf and the Emin Pasha Gulf, whereas Haplochromis goldschmidti sp. n. was only found in the Emin Pasha Gulf. Of the latter species only males are available, but it seems unlikely that it represents a case of male colour polymorphism as several presumably unrelated characters differ in sympatry between the two species, suggesting that there is no gene flow. Statistical analysis revealed that the overall difference between the two species is greater than that between the populations from the two locations. Body depth of the two species in sympatry in the Emin Pasha Gulf was more similar than that of Haplochromis goldschmidti sp. n. and the allopatric population of Haplochromis argens sp. n. from the Mwanza Gulf,which mayindicate an overall environmental effect. However, several measurements related to the width of snout and mouth differed more between the populations of the two species in sympatry than between the allopatric populations. In contrast to a group of zooplanktivorous species that recovered successfully after environmental changes in the lake, Haplochromis argens sp. n. is among a group that became extremely rare and probably is in danger of extinction; the conservation status of Haplochromis goldschmidti sp. n. is currently unknown. PMID:23717179
Kautt, Andreas F; Machado-Schiaffino, Gonzalo; Meyer, Axel
The formation of species in the absence of geographic barriers (i.e. sympatric speciation) remains one of the most controversial topics in evolutionary biology. While theoretical models have shown that this most extreme case of primary divergence-with-gene-flow is possible, only a handful of accepted empirical examples exist. And even for the most convincing examples uncertainties remain; complex histories of isolation and secondary contact can make species falsely appear to have originated by sympatric speciation. This alternative scenario is notoriously difficult to rule out. Midas cichlids inhabiting small and remote crater lakes in Nicaragua are traditionally considered to be one of the best examples of sympatric speciation and lend themselves to test the different evolutionary scenarios that could lead to apparent sympatric speciation since the system is relatively small and the source populations known. Here we reconstruct the evolutionary history of two small-scale radiations of Midas cichlids inhabiting crater lakes Apoyo and Xiloá through a comprehensive genomic data set. We find no signs of differential admixture of any of the sympatric species in the respective radiations. Together with coalescent simulations of different demographic models our results support a scenario of speciation that was initiated in sympatry and does not result from secondary contact of already partly diverged populations. Furthermore, several species seem to have diverged simultaneously, making Midas cichlids an empirical example of multispecies outcomes of sympatric speciation. Importantly, however, the demographic models strongly support an admixture event from the source population into both crater lakes shortly before the onset of the radiations within the lakes. This opens the possibility that the formation of reproductive barriers involved in sympatric speciation was facilitated by genetic variants that evolved in a period of isolation between the initial founding
Kautt, Andreas F.; Machado-Schiaffino, Gonzalo; Meyer, Axel
The formation of species in the absence of geographic barriers (i.e. sympatric speciation) remains one of the most controversial topics in evolutionary biology. While theoretical models have shown that this most extreme case of primary divergence-with-gene-flow is possible, only a handful of accepted empirical examples exist. And even for the most convincing examples uncertainties remain; complex histories of isolation and secondary contact can make species falsely appear to have originated by sympatric speciation. This alternative scenario is notoriously difficult to rule out. Midas cichlids inhabiting small and remote crater lakes in Nicaragua are traditionally considered to be one of the best examples of sympatric speciation and lend themselves to test the different evolutionary scenarios that could lead to apparent sympatric speciation since the system is relatively small and the source populations known. Here we reconstruct the evolutionary history of two small-scale radiations of Midas cichlids inhabiting crater lakes Apoyo and Xiloá through a comprehensive genomic data set. We find no signs of differential admixture of any of the sympatric species in the respective radiations. Together with coalescent simulations of different demographic models our results support a scenario of speciation that was initiated in sympatry and does not result from secondary contact of already partly diverged populations. Furthermore, several species seem to have diverged simultaneously, making Midas cichlids an empirical example of multispecies outcomes of sympatric speciation. Importantly, however, the demographic models strongly support an admixture event from the source population into both crater lakes shortly before the onset of the radiations within the lakes. This opens the possibility that the formation of reproductive barriers involved in sympatric speciation was facilitated by genetic variants that evolved in a period of isolation between the initial founding
Cohen, Andrew S.; Blome, Margaret; Ivory, Sarah; King, John; Cole, Julie; McGlue, Michael
Long records of Quaternary ecological and climatic change are critical to understanding the range of potential responses of ecosystems to environmental forcing. Here we present an integrated lake and watershed paleoecological analysis from drill core records obtained by the Lake Malawi Drilling Project, documenting extraordinary fluctuations in climate, hydrology and ecosystem response for the southern tropics of Africa. High resolution lacustrine and terrestrial paleoecology and sedimentology data sets from these Early Pleistocene-Holocene drill cores provide the most complete record of this duration currently available from Africa. Time series analyses of these records demonstrate strong orbital forcing of regional hydroclimate that drives high-amplitude changes in Malawi ecosystems. Prior to ~600ka we also observe a secondary overprint of watershed processes involving river capture or diversion that may have a tectonic origin. We observe shifts between more arid conditions (shallow alkaline and well mixed lake, with discontinuous desert vegetation) and more humid environments (deep, stratified, freshwater lake with dense forest). These broadly synchronous changes in lake paleoecology, lake sedimentology, and watershed vegetation demonstrate the major role of climate in regulating this system. Transitions between these lake/watershed state extremes is often very abrupt, suggesting that the combined lake/watershed repeatedly passed through hydroclimate thresholds, with important implications for the evolution of the lake's endemic biodiversity and ecosystem. The tempo of lake/watershed state fluctuations changes at the Mid-Pleistocene Transition, altering from one of higher frequency/lower amplitude variability prior to 900ka to lower frequency/higher amplitude variability after that time.
Svensson, Ola; Woodhouse, Katie; van Oosterhout, Cock; Smith, Alan; Turner, George F; Seehausen, Ole
The genetic architecture of mate preferences is likely to affect significant evolutionary processes, including speciation and hybridization. Here, we investigate laboratory hybrids between a pair of sympatric Lake Victoria cichlid fish species that appear to have recently evolved from a hybrid population between similar predecessor species. The species demonstrate strong assortative mating in the laboratory, associated with divergent male breeding coloration (red dorsum versus blue). We show in a common garden experiment, using DNA-based paternity testing, that the strong female mate preferences among males of the two species are fully recovered in a large fraction of their F2 hybrid generation. Individual hybrid females often demonstrated consistent preferences in multiple mate choice trials (more than or equal to five) across a year or more. This result suggests that female mate preference is influenced by relatively few major genes or genomic regions. These preferences were not changed by experience of a successful spawning event with a male of the non-preferred species in a no-choice single-male trial. We found no evidence for imprinting in the F2 hybrids, although the F1 hybrid females may have been imprinted on their mothers. We discuss this nearly Mendelian inheritance of consistent innate mate preferences in the context of speciation theory.
Katoh, Rei; Munehara, Hiroyuki; Kohda, Masanori
Telmatochromis temporalis is a bi-parental substrate brooding cichlid endemic to Lake Tanganyika. Paired males were always larger than their mates and had territories around nests against conspecific males. However, males smaller than the paired females were found in 18% of the nests. Here we report a reproductive tactic of these small males. The small males had as heavy gonads as paired males, and the gonad somatic index (GSI) of the small males was much higher than that of the latter. The examinations of the paternity and maternity using microsatellite-DNA as a genetic marker revealed that the small males were not genetically related to the pair members, and sired some young in 3 of 5 nests. These small males did not guard the broods, suggesting that they are likely to perform reproductive parasitism as sneakers. Paired males could not enter their spawning nests due to their large size, which made it difficult to chase out sneakers once they entered the nest. Some males as small as the sneakers were found outside the territories of paired males, and their gonads were quite small. Circumstantial evidence suggests that small males have two alternative investment patterns: investing in gonad to be sneakers, and investing in growth to probably be territorial males.
Imanishi's "mental" (cerebral) view of speciation is presented, in Mizuhata's revision. The key concept here is the "ethological partition" of the species. Members of each species=society (etho-species) share the same mental (brain) software, irrespective of their genetic structure. Cerebral animals perform active programmed selection, not to be confused with passive, non-programmed "natural selection" as in Neo-Darwinism. The program includes mating-choice of peculiar characters, distinct from the Neo-Darwinian sexual selection supposed due to the specific choosy genes. Speciation can occur, as a "partition of species=society", with bifurcation of mate-choosing program in the parent species. A main promoter for this bifurcation is species-specific "passion" for especially significant characters: long necks, ornamental antlers, ocelli feathers, bright nuptial colors etc. The cichlids in Lake Victoria achieved explosive speciation, while retaining their genetic homogeneity completely. Therefore it is illogical to attribute this divergence to extraordinary mutations in "action controlling genes". The origin of species=society (etho-species) can trace along to the Cambrian Period.
Hayward, A; Tsuboi, M; Owusu, C; Kotrschal, A; Buechel, S D; Zidar, J; Cornwallis, C K; Løvlie, H; Kolm, N
Parasite diversity and abundance (parasite load) vary greatly among host species. However, the influence of host traits on variation in parasitism remains poorly understood. Comparative studies of parasite load have largely examined measures of parasite species richness and are predominantly based on records obtained from published data. Consequently, little is known about the relationships between host traits and other aspects of parasite load, such as parasite abundance, prevalence and aggregation. Meanwhile, understanding of parasite species richness may be clouded by limitations associated with data collation from multiple independent sources. We conducted a field study of Lake Tanganyika cichlid fishes and their helminth parasites. Using a Bayesian phylogenetic comparative framework, we tested evolutionary associations between five key host traits (body size, gut length, diet breadth, habitat complexity and number of sympatric hosts) predicted to influence parasitism, together with multiple measures of parasite load. We find that the number of host species that a particular host may encounter due to its habitat preferences emerges as a factor of general importance for parasite diversity, abundance and prevalence, but not parasite aggregation. In contrast, body size and gut size are positively related to aspects of parasite load within, but not between species. The influence of host phylogeny varies considerably among measures of parasite load, with the greatest influence exerted on parasite diversity. These results reveal that both host morphology and biotic interactions are key determinants of host-parasite associations and that consideration of multiple aspects of parasite load is required to fully understand patterns in parasitism.
Kavembe, Geraldine D; Kautt, Andreas F; Machado-Schiaffino, Gonzalo; Meyer, Axel
Ecological diversification through divergent selection is thought to be a major force during the process of adaptive radiations. However, the large sizes and complexity of most radiations such as those of the cichlids in the African Great Lakes make it impossible to infer the exact evolutionary history of any population divergence event. The genus Alcolapia, a small cichlid lineage endemic to Lakes Magadi and Natron in East Africa, exhibits phenotypes similar to some of those found in cichlids of the radiations of the African Great Lakes. The simplicity within Alcolapia makes it an excellent model system to investigate ecological diversification and speciation. We used an integrated approach including population genomics based on RAD-seq data, geometric morphometrics and stable isotope analyses to investigate the eco-morphological diversification of tilapia in Lake Magadi and its satellite lake Little Magadi. Additionally, we reconstructed the demographic history of the species using coalescent simulations based on the joint site frequency spectrum. The population in Little Magadi has a characteristically upturned mouth--possibly an adaptation to feeding on prey from the water surface. Eco-morphological differences between populations within Lake Magadi are more subtle, but are consistent with known ecological differences between its lagoons such as high concentrations of nitrogen attributable to extensive guano deposits in Rest of Magadi relative to Fish Springs Lagoon. All populations diverged simultaneously only about 1100 generations ago. Differences in levels of gene flow between populations and the effective population sizes have likely resulted in the inferred heterogeneous patterns of genome-wide differentiation.
Dalton, Brian E; Cronin, Thomas W; Marshall, N Justin; Carleton, Karen L
The extent of animal colouration is determined by an interplay between natural and sexual selection. Both forces probably shape colouration in the speciose, rock-dwelling cichlids of Lake Malawi. Sexual selection is thought to drive male colouration, overcoming natural selection to create conspicuous colour patterns via female mate choice and male-male competition. However, natural selection should make female cichlids cryptic because they mouthbrood their young. We hypothesize that as a result of both sexual and natural selection, males will have colours that are more conspicuous than female colours. Cichlid spectral sensitivity, especially in the ultraviolet, probably influences how colours appear to them. Here we use simple models of the trichromatic colour space of cichlid visual systems to compare the conspicuousness of male and female nuptial colours of nine species. Conspicuousness of colours was evaluated as their Euclidian distance in colour space from environmental backgrounds and from other colours on the same fish. We find in six of the nine species that breeding males have colours that are statistically more conspicuous than female colours. These colours contrast strongly with each other or with the backgrounds, and they fall within a range of spectra best transmitted in the habitat. Female colour distances were sometimes smaller, suggesting that females of some species are more cryptic than males. Therefore, selection can differentially act to generate male colours that are more conspicuous than those in females. However, in two species, females had colours that were more conspicuous than male colours, suggesting that other selective forces and possibly sexual conflicts are acting in this system.
Wright, D S; Demandt, N; Alkema, J T; Seehausen, O; Groothuis, T G G; Maan, M E
Local adaptation can be a potent force in speciation, with environmental heterogeneity leading to niche specialization and population divergence. However, local adaption often requires nonrandom mating to generate reproductive isolation. Population divergence in sensory properties can be particularly consequential in speciation, affecting both ecological adaptation and sexual communication. Pundamilia pundamila and Pundamilia nyererei are two closely related African cichlid species that differ in male coloration, blue vs. red. They co-occur at rocky islands in southern Lake Victoria, but inhabit different depth ranges with different light environments. The species differ in colour vision properties, and females exert species-specific preferences for blue vs. red males. Here, we investigated the mechanistic link between colour vision and preference, which could provide a rapid route to reproductive isolation. We tested the behavioural components of this link by experimentally manipulating colour perception - we raised both species and their hybrids under light conditions mimicking shallow and deep habitats - and tested female preference for blue and red males under both conditions. We found that rearing light significantly affected female preference: shallow-reared females responded more strongly to P. pundamilia males and deep-reared females favoured P. nyererei males - implying that visual development causally affects mate choice. These results are consistent with sensory drive predictions, suggesting that the visual environment is key to behavioural isolation of these species. However, the observed plasticity could also make the species barrier vulnerable to environmental change: species-assortative preferences were weaker in females that were reared in the other species' light condition.
Scholz, C. A.; Johnson, T. C.; King, J.; Cohen, A. S.; Lyons, R. P.; Kalindekafe, L.; Forman, S. L.; McHargue, L. R.; Singer, B. S.
Lake Malawi is one of the largest and oldest lakes in the southern hemisphere and in the tropical latitudes, measuring more than 580 km in length, with a maximum depth of 700 m, and with an estimated age of more than 7 Ma. The lake water column is anoxic below 200 m, thus preserving an extensive record of laminated (varved) sediments. The lake's hydrologic budget is hypersensitive to minor changes in precipitation:evaporation, and signals of lake level and limnologic change respond markedly to regional climate variations. During February-March 2005 a scientific drilling program was conducted on the deep, open waters of Lake Malawi, using a converted local fuel barge outfitted with a deep-water-capable geotechnical drilling rig, a portable dynamic positioning system, and a suite of sampling tools designed for deep lake coring. A total of 623 m of core was recovered in seven holes at two sites on the lake. The high-resolution North Basin site (water depth 359 m) was triple-cored, providing redundant sampling of the sedimentary section representing the past ~75-100 kyr; the Central Basin deep site (592 m water depth) was double cored over the period of the past ~200 kyr, and single-cored to 380 m, covering perhaps 1.5 million years. The North Basin high-resolution site targeted a hemipelagic sequence, which in core samples consists of alternating zones of laminated and homogenous silty mud. At the base of these cores we observe well-sorted medium-grained shoreface sand deposits, that we interpret to be a transgressive surface deposited following a severe (500 m) lake lowstand. Density, magnetic susceptibility, TOC, and C/N profiles closely follow lithologic changes, and correspond to periods of enhanced or diminished productivity and dramatically varying lake level. Initial dating of the North Basin site (total core length 38 m) is underway using a combination of Radiocarbon, Tephra Ar-Ar, and Optically-Stimulated Luminescence methods, and initial results indicate a
Navon, Dina; Olearczyk, Nathan; Albertson, R Craig
Adaptive radiations are often characterized by the rapid evolution of traits associated with divergent feeding modes. For example, the evolutionary history of African cichlids is marked by repeated and coordinated shifts in skull, trophic, fin and body shape. Here, we seek to explore the molecular basis for fin shape variation in Lake Malawi cichlids. We first described variation within an F2 mapping population derived by crossing two cichlid species with divergent morphologies including fin shape. We then used this population to genetically map loci that influence variation in this trait. We found that the genotype-phenotype map for fin shape is largely distinct from other morphological characters including body and craniofacial shape. These data suggest that key aspects of fin, body and jaw shape are genetically modular and that the coordinated evolution of these traits in cichlids is more likely due to common selective pressures than to pleiotropy or linkage. We next combined genetic mapping data with population-level genome scans to identify wnt7aa and col1a1 as candidate genes underlying variation in the number of pectoral fin ray elements. Gene expression patterns across species with different fin morphologies and small molecule manipulation of the Wnt pathway during fin development further support the hypothesis that variation at these loci underlies divergence in fin shape between cichlid species. In all, our data provide additional insights into the genetic and molecular mechanisms associated with morphological divergence in this important adaptive radiation.
Albertson, R Craig; Kocher, Thomas D
The role of hybridization in the evolution of animal species is poorly understood. Transgressive segregation is a mechanism through which hybridization can generate diversity and ultimately lead to speciation. In this report we investigated the capacity of hybridization to generate novel (transgressive) phenotypes in the taxonomically diverse cichlid fishes. We generated a large F2 hybrid population by crossing two closely related cichlid species from Lake Malawi in Africa with differently shaped heads. Our morphometric analysis focused on two traits with different selective histories. The cichlid lower jaw (mandible) has evolved in response to strong directional selection, and does not segregate beyond the parental phenotype. The cichlid neurocranium (skull) has likely diverged in response to forces other than consistent directional selection (e.g., stabilizing selection), and exhibits marked transgressive segregation in our F2 population. We show that the genetic architecture of the cichlid jaw limits transgression, whereas the genetic basis of skull shape is permissive of transgressive segregation. These data suggest that natural selection, acting through the genome, will limit the degree of diversity that may be achieved via hybridization. Results are discussed in the context of the broader question of how phenotypic diversity may be achieved in rapidly evolving systems.
Shillington, D. J.; Scholz, C. A.; Gaherty, J. B.; Accardo, N. J.; McCartney, T.; Chindandali, P. R. N.; Kamihanda, G.; Trinhammer, P.; Wood, D. A.; Khalfan, M.; Ebinger, C. J.; Nyblade, A.; Mbogoni, G. J.; Mruma, A. H.; Salima, J.; Ferdinand-Wambura, R.
Little is known about the controls on the initiation and development of magmatism and segmentation in young rift systems. The northern Lake Malawi (Nyasa) rift in the East African Rift System is an early stage rift exhibiting pronounced tectonic segmentation, which is defined in the upper crust by ~100-km-long border faults. Very little volcanism is associated with rifting; the only surface expression of magmatism occurs in an accommodation zone between segments to the north of the lake in the Rungwe Volcanic Province. The SEGMeNT (Study of Extension and maGmatism in Malawi aNd Tanzania) project is a multidisciplinary, multinational study that is acquiring a suite of geophysical, geological and geochemical data to characterize deformation and magmatism in the crust and mantle lithosphere along 2-3 segments of this rift. As a part of the SEGMeNT project, we acquired seismic reflection and refraction data in Lake Malawi (Nyasa) in March-April 2015. Over 2000 km of seismic reflection data were acquired with a 500 to 2580 cu in air gun array from GEUS/Aarhus and a 500- to 1500-m-long seismic streamer from Syracuse University over a grid of lines across and along the northern and central basins. Air gun shots from MCS profiles and 1000 km of additional shooting with large shot intervals were also recorded on 27 short-period and 6 broadband lake bottom seismometers from Scripps Oceanographic Institute as a part of the Ocean Bottom Seismic Instrument Pool (OBSIP) as well as the 55-station onshore seismic array. The OBS were deployed along one long strike line and two dip lines. We will present preliminary data and results from seismic reflection and refraction data acquired in the lake and their implications for crustal deformation within and between rift segments. Seismic reflection data image structures up to ~5-6 km below the lake bottom, including syntectonic sediments, intrabasinal faults and other complex horsts. Some intrabasinal faults in both the northern and
Geophysical data are currently being interpreted as evidence for a late Pleistocene desiccation of Lake Victoria and its refilling 14,600 years ago. This implies that between 500 and 1000 endemic cichlid fish species must have evolved in 14,600 years, the fastest large-scale species radiation known. A recent review concludes that biological evidence clearly rejects the postulated Pleistocene desiccation of the lake: a 14,600 year history would imply exceptionally high speciation rates across a range of unrelated fish taxa. To test this suggestion, I calculated speciation rates for all 41 phylogenetic lineages of fish in the lake. Except for one cichlid lineage, accepting a 14 600 year history does not require any speciation rates that fall outside the range observed in fishes in other young lakes around the world. The exceptional taxon is a lineage of haplochromine cichlids that is also known for its rapid speciation elsewhere. Moreover, since it is unknown how many founding species it has, it is not certain that its speciation rates are really outside the range observed in fishes in other young lakes. Fish speciation rates are generally faster in younger than in older lakes, and those in Lake Victoria, by far the largest of the young lakes of the world, are no exception. From the speciation rates and from biogeographical observations that Lake Victoria endemics, which lack close relatives within the lake basin, have such relatives in adjacent drainage systems that may have had Holocene connections to Lake Victoria, I conclude that the composition of the fish assemblage does not provide biological evidence against Pleistocene desiccation. It supports a hypothesis of recent colonization from outside the lake basin rather than survival of a diverse assemblage within the basin. PMID:11886641
Martin, Christopher H; Cutler, Joseph S; Friel, John P; Dening Touokong, Cyrille; Coop, Graham; Wainwright, Peter C
One of the most celebrated examples of sympatric speciation in nature are monophyletic radiations of cichlid fishes endemic to Cameroon crater lakes. However, phylogenetic inference of monophyly may not detect complex colonization histories involving some allopatric isolation, such as double invasions obscured by genome-wide gene flow. Population genomic approaches are better suited to test hypotheses of sympatric speciation in these cases. Here, we use comprehensive sampling from all four sympatric crater lake cichlid radiations in Cameroon and outgroups across Africa combined with next-generation sequencing to genotype tens of thousands of SNPs. We find considerable evidence of gene flow between all four radiations and neighboring riverine populations after initial colonization. In a few cases, some sympatric species are more closely related to outgroups than others, consistent with secondary gene flow facilitating their speciation. Our results do not rule out sympatric speciation in Cameroon cichlids, but rather reveal a complex history of speciation with gene flow, including allopatric and sympatric phases, resulting in both reproductively isolated species and incipient species complexes. The best remaining non-cichlid examples of sympatric speciation all involve assortative mating within microhabitats. We speculate that this feature may be necessary to complete the process of sympatric speciation in nature.
Wagner, Catherine E; Keller, Irene; Wittwer, Samuel; Selz, Oliver M; Mwaiko, Salome; Greuter, Lucie; Sivasundar, Arjun; Seehausen, Ole
Although population genomic studies using next generation sequencing (NGS) data are becoming increasingly common, studies focusing on phylogenetic inference using these data are in their infancy. Here, we use NGS data generated from reduced representation genomic libraries of restriction-site-associated DNA (RAD) markers to infer phylogenetic relationships among 16 species of cichlid fishes from a single rocky island community within Lake Victoria's cichlid adaptive radiation. Previous attempts at sequence-based phylogenetic analyses in Victoria cichlids have shown extensive sharing of genetic variation among species and no resolution of species or higher-level relationships. These patterns have generally been attributed to the very recent origin (<15,000 years) of the radiation, and ongoing hybridization between species. We show that as we increase the amount of sequence data used in phylogenetic analyses, we produce phylogenetic trees with unprecedented resolution for this group. In trees derived from our largest data supermatrices (3 to >5.8 million base pairs in width), species are reciprocally monophyletic with high bootstrap support, and the majority of internal branches on the tree have high support. Given the difficulty of the phylogenetic problem that the Lake Victoria cichlid adaptive radiation represents, these results are striking. The strict interpretation of the topologies we present here warrants caution because many questions remain about phylogenetic inference with very large genomic data set and because we can with the current analysis not distinguish between effects of shared ancestry and post-speciation gene flow. However, these results provide the first conclusive evidence for the monophyly of species in the Lake Victoria cichlid radiation and demonstrate the power that NGS data sets hold to resolve even the most difficult of phylogenetic challenges.
Oyugi, Dalmas Oyugi; Harper, David Matthew; Ntiba, Japheth Michemi; Kisia, Seth Marande; Britton, John Robert
The tilapiine cichlids Oreochromis leucostictus and Tilapia zillii were introduced into Lake Naivasha, Kenya, in 1956. Previous studies on data collected to 1987 revealed they were persistent following establishment, despite environmental variability and exploitation. Recent data, however, suggest this persistence is under threat as data indicate some significant declines in aspects of their abundance since 1999. The influence of changes in lake level, allodiversity and fishing effort on this decline was tested and showed that a decline in lake level was a significant causal factor. The recent change in allodiversity, with the establishment and dominance of Cyprinus carpio in the fishery, was not significant on the catch per unit effort of O. leucostictus but was on T. zillii. Since 1999, catches of tilapiines in the fishery have been independent of fishing effort, contrary to between 1975 and 1987, suggesting their management through application of fishery models may no longer be applicable. As it was anthropogenic-mediated lake level changes that were mainly responsible for their decline, then lake management should focus on sustainable water utilization that maximizes lake levels in accordance with the basin-wide water balance.
Castañeda, I. S.; Werne, J. P.; Johnson, T. C.
In general, information regarding tropical African climate variability is relatively limited, especially in comparison with high-latitude studies. Unlike the high-latitudes where climate change is often expressed by fluctuations in temperature, low-latitude climate change is often expressed as variability in zonal circulation, which can result in hydrological fluctuations. Lake Malawi, situated in low-latitude tropical Africa (9-14° S), contains a continuous and high-resolution sedimentary record of the past 22ka BP and is anoxic below 250m, which enhances preservation of organic matter (OM). For these reasons, L. Malawi is an excellent location to examine the response of low-latitude African climate to global climate change. The climate of Malawi is strongly influenced by the position and seasonal migration of the ITCZ. During the rainy season from November to March, the ITCZ is positioned over L. Malawi (12-13° S) and the dominant winds are weak and northerly. Between April and May the ITCZ moves northward towards the equator and strong southerly winds prevail (Jury & Mwafulirwa, 2002). Previous studies of L. Malawi have shown responses to global climatic events, such as the Younger Dryas. Additionally, studies have demonstrated the response of L. Malawi to local or regional events, such as variability in the ITCZ. Based on BSi MAR, diatom, phosphorus, and trace metal data, Johnson et al. (2002) proposed that at times more frequent or stronger northerly winds promoted upwelling in the northern basin of L. Malawi, and suggested more southerly migrations of the ITCZ (reaching latitudes of >13\\deg S) as the cause of these increased winds. Additionally, a recent study of L. Malawi based on multiple bulk geochemical proxies provides evidence for both southward and northward displacements of the ITCZ during the past 23ka BP (Filippi and Talbot, submitted). In this study the molecular biomarker record of L. Malawi is examined. Previous studies of Lake Malawi have
In the rare perennial bodies of water of the Sahara desert, only a few fish species have survived to increasing aridification since the end of the last humid period at the Holocene, approximately 5000 years BP. Here, I report the occurrence of an undescribed haplochomine cichlid fish in Lake Boukou, one of the seven Ounianga Serir lakes (Chad). These lakes are located in one of the most arid areas of the Sahara desert, but they persist by virtue of subsurface inflow of fresh groundwater from a large fossil aquifer. Astatotilapia tchadensis sp. nov. is characterized by a black bar between eye and corner of mouth, rounded orange spots on anal fin, scales ctenoid, lower limb of first gill arch with 7-8 gill rackers, dorsal fin with 13-14 spines and 9-11 soft rays, anal fin with 3 spines and 8-9 soft rays, 29 or 30 lateral line scales, and lower pharyngeal dentition with enlarged molariform teeth. The new species is easily distinguished from A. desfontainii and A. flaviijosephii, the northernmost haplochromine species currently isolated from its other group members, and appears close to an unnamed species of Lake Chad basin. Ounianga Serir lakes and especially Lake Boukou present a remarkable diversity of fish, the highest known in the Sahara desert with a total of at least six fish species belonging to six genera and three families. They also constitute an exceptional natural landscape inscribed on the UNESCO world heritage list in 2012 and a biodiversity hotspot for desert vertebrate species.
Ivory, S.; Lézine, A. M.; Vincens, A.; Cohen, A. S.
African ecosystems are at great risk due to climate and land-use change. Despite the status of several of these regions as biodiversity hotspots, long-standing ideas about African ecology and biogeography have been unable to be tested until now due to lack of sufficiently long records. Here, we present the first long, continuous terrestrial record of vegetation from Lake Malawi, East Africa which goes back to the early Late Quaternary, permitting us to investigate changes in physiognomy and forest composition over many transitions. In this record, we observe eight phases of forest expansion and collapse. Although diversity is much greater during forest phases, composition varies little from phase to phase. Very high abundances of afromontane taxa suggest frequent widespread colonization of the lowlands by modern high elevation trees. Although there are clear successional stages within each forest such that turnover is great within a single phase, among forest samples between phases, there is little dissimilarity. Each forest phase is interrupted by rapid decline of arboreal taxa and expansion of semi-arid grasslands or woodlands whose composition varies greatly from phase to phase. The variable composition of the more open phases, all occurring during arid periods, is likely dynamically linked to thresholds in regional hydrology associated with lake level and moisture recycling within the watershed. This vegetation is unlike any found at Malawi today, with assemblages suggesting strong Somali-Masai affinities. Furthermore, nearly all semi-arid assemblages contain small abundances of forest taxa typically growing in areas with wetter edaphic conditions, suggesting that moist lowland gallery forests were present but restricted to waterways during exceptionally arid times. The waxing and waning of forests throughout this interval has important implications for early human biogeography across Africa as well as disturbance regimes that are crucial for the maintenance of
Parsons, Kevin J.; Cooper, W. James; Albertson, R. Craig
The African cichlids of the East-African rift-lakes provide one of the most dramatic examples of adaptive radiation known. It has long been thought that functional decoupling of the oral and pharyngeal jaws in cichlids has facilitated their explosive evolution. Recent research has also shown that craniofacial evolution from radiations in lakes Victoria, Malawi, and Tanganyika has occurred along a shared primary axis of shape divergence, whereby the preorbital region of the skull changes in a manner that is, relatively independent from other head regions. We predicted that the preorbital region would comprise a variational module and used an extensive dataset from each lake that allowed us to test this prediction using a model selection approach. Our findings supported the presence of a preorbital module across all lakes, within each lake, and for Malawi, within sand and rock-dwelling clades. However, while a preorbital module was consistently present, notable differences were also observed among groups. Of particular interest, a negative association between patterns of variational modularity was observed between the sand and rock-dwelling clades, a patter consistent with character displacement. These findings provide the basis for further experimental research involving the determination of the developmental and genetic bases of these patterns of modularity. PMID:21716745
High-resolution, air-gun-sourced seismic reflection surveys over the offshore regions of five river deltas in Lake Malawi in the East African rift system reveal considerable variability in acoustic facies and stratigraphic architecture. This variability can largely be attributed to the influences of different structural settings, and to a lesser degree to high-amplitude (100-400 m) and high-frequency (1000 to 100,000 yr) fluctuations in lake level. Deltas on flexural and axial margins in the rift lake show well-developed progradational geometries. In contrast, a delta on a steep, accommodation zone margin distributes coarse sediments over a broad depositional apron, rather than concentrating sediment in discrete progradational lobes as on the other deltas. A large border fault margin river delta displays the most complex tectonic and stratigraphic architecture of all the deltas studied. It contains several delta-associated facies, including prograding clinoform packages, fan deltas stacked against a boundary fault, and extensive subaqueous fans. Flexural margin lowstand deltas may be the most prospective for hydrocarbon exploration due to their large, internally well-organized, progradational lobes and their close proximity to deep-water, high total organic carbon lacustrine source facies.
Keller, I; Wagner, C E; Greuter, L; Mwaiko, S; Selz, O M; Sivasundar, A; Wittwer, S; Seehausen, O
Adaptive radiations are an important source of biodiversity and are often characterized by many speciation events in very short succession. It has been proposed that the high speciation rates in these radiations may be fuelled by novel genetic combinations produced in episodes of hybridization among the young species. The role of such hybridization events in the evolutionary history of a group can be investigated by comparing the genealogical relationships inferred from different subsets of loci, but such studies have thus far often been hampered by shallow genetic divergences, especially in young adaptive radiations, and the lack of genome-scale molecular data. Here, we use a genome-wide sampling of SNPs identified within restriction site-associated DNA (RAD) tags to investigate the genomic consistency of patterns of shared ancestry and adaptive divergence among five sympatric cichlid species of two genera, Pundamilia and Mbipia, which form part of the massive adaptive radiation of cichlids in the East African Lake Victoria. Species pairs differ along several axes: male nuptial colouration, feeding ecology, depth distribution, as well as the morphological traits that distinguish the two genera and more subtle morphological differences. Using outlier scan approaches, we identify signals of divergent selection between all species pairs with a number of loci showing parallel patterns in replicated contrasts either between genera or between male colour types. We then create SNP subsets that we expect to be characterized to different extents by selection history and neutral processes and describe phylogenetic and population genetic patterns across these subsets. These analyses reveal very different evolutionary histories for different regions of the genome. To explain these results, we propose at least two intergeneric hybridization events (between Mbipia spp. and Pundamilia spp.) in the evolutionary history of these five species that would have lead to the evolution
Lane, Christine S; Chorn, Ben T; Johnson, Thomas C
The most explosive volcanic event of the Quaternary was the eruption of Mt. Toba, Sumatra, 75,000 y ago, which produced voluminous ash deposits found across much of the Indian Ocean, Indian Peninsula, and South China Sea. A major climatic downturn observed within the Greenland ice cores has been attributed to the cooling effects of the ash and aerosols ejected during the eruption of the Youngest Toba Tuff (YTT). These events coincided roughly with a hypothesized human genetic bottleneck, when the number of our species in Africa may have been reduced to near extinction. Some have speculated that the demise of early modern humans at that time was due in part to a dramatic climate shift triggered by the supereruption. Others have argued that environmental conditions would not have been so severe to have such an impact on our ancestors, and furthermore, that modern humans may have already expanded beyond Africa by this time. We report an observation of the YTT in Africa, recovered as a cryptotephra layer in Lake Malawi sediments, >7,000 km west of the source volcano. The YTT isochron provides an accurate and precise age estimate for the Lake Malawi paleoclimate record, which revises the chronology of past climatic events in East Africa. The YTT in Lake Malawi is not accompanied by a major change in sediment composition or evidence for substantial temperature change, implying that the eruption did not significantly impact the climate of East Africa and was not the cause of a human genetic bottleneck at that time.
Takeuchi, Yuichi; Hori, Michio; Oda, Yoichi
Behavioral lateralization has been documented in many vertebrates. The scale-eating cichlid fish Perissodus microlepis is well known for exhibiting lateral dimorphism in its mouth morphology and lateralized behavior in robbing scales from prey fish. A previous field study indicated that this mouth asymmetry closely correlates with the side on which prey is attacked, but details of this species' predation behavior have not been previously analyzed because of the rapidity of the movements. Here, we studied scale-eating behavior in cichlids in a tank through high-speed video monitoring and quantitative assessment of behavioral laterality and kinematics. The fish observed showed a clear bias toward striking on one side, which closely correlated with their asymmetric mouth morphologies. Furthermore, the maximum angular velocity and amplitude of body flexion were significantly larger during attacks on the preferred side compared to those on the nonpreferred side, permitting increased predation success. In contrast, no such lateral difference in movement elements was observed in acoustically evoked flexion during the escape response, which is similar to flexion during scale eating and suggests that they share a common motor control pathway. Thus the neuronal circuits controlling body flexion during scale eating may be functionally lateralized upstream of this common motor pathway. PMID:22238598
Johnson, T. C.; Powers, L. A.; Werne, J. P.; Brown, E. T.; Castaneda, I.; Schouten, S.; Sinninghe-Damste, J.
Piston and multi-cores recovered from the north basin of Lake Malawi in 1998 by the International Decade for the East African Lakes (IDEAL) have provided a rich history of climate variability spanning the past 25,000 years. As we now begin to analyze the cores recovered by the Malawi Drilling Project in early 2005, we are considering the relationships among sedimentary signals of temperature (TEX86), northerly winds associated with a southward excursion of the Inter-Tropical Convergence Zone (per cent biogenic silica), and rainfall (terrigenous mass accumulation rate) in the well dated 1998 cores. A high-resolution record of the past 800 years suggests that rainfall in this region (10 - 12° S, 30 - 35° E) was relatively low during the Little Ice Age, when northerly winds were more prevalent, attributed to a more southerly position of the ITCZ during austral summers. The TEX86 signal of lake (surface?) temperature ranged mostly between 24 and 26°C during this period, with the coldest temperature of about 22°C around AD1680 and the warmest temperature, exceeding 27°C, in the youngest sediment sample. The cooler water temperatures coincide with periods of highest diatom productivity, consistent with the latter being due to relatively intense upwelling associated with the northerly winds. Our observation of low rainfall during periods of more southerly migration of the ITCZ is consistent with the results of McHugh and Rogers (2001), who linked rainfall in southeastern Africa to the North Atlantic Oscillation (NAO). During years of weak NAO, equatorial westerly transport of Atlantic moisture across Africa during austral summer is relatively intense, causing high rainfall in the East African Rift between the equator and 16° S. Conversely, when the NAO is positive, rainfall is higher south of 15° S than north of this latitude, which is consistent with a southward migration of the ITCZ. McHugh, M. J. and J. C. Rogers (2001). "North Atlantic Oscillation influence on
Brown, E. T.; Johnson, T. C.; Scholz, C. A.; King, J. W.; Cohen, A. S.
As part of the Lake Malawi Drilling Project we have undertaken a 1 cm-scale resolution (ca. 20 yr) XRF scanning study of core from site 2A in the Northern Basin of the lake. We have focused on certain parameters, including Fe, Zr:Ti, and Si:Ti; these may be interpreted to represent changing input of terrigenous sediments (delivered to the lake by rivers), soils developed from weathered volcanic ash (delivered more effectively under drier windier conditions), and biogenic silica, respectively. We present here a record of East African climate since 55 ka that indicates rapid shifts between discrete climate modes that appear to be related to abrupt warming (Dansgaard- Oeschger) and cooling (Heinrich) events observed in Greenland. The events in Malawi appear to correspond to episodes of dry conditions and increased northerly winds. Such conditions may occur at times that southward excursion of the Intertropical Convergence Zone brings the tropical rain belt to the south of the study area. Although the timing of the Malawi events cannot be determined exactly, our independent age model, based on 25 radiocarbon dates and stratigraphic correlations to events dated in a nearby at 63 and 75 ka, implies that they occur prior to their Greenland counterparts, consistent with southward excursions of the ITCZ during Greenland stadials. The magnitude and form of each of the events recorded in Malawi sediments corresponds to the scale of the subsequent Greenland warming. This suggests that a tropical component of climate sets a template for abrupt high northern latitude climate fluctuations associated with the bipolar seesaw.
Concannon, Moira R; Albertson, R Craig
The evolution of an exaggerated trait can lead to a novel morphology that allows organisms to exploit new niches. The molecular bases of such phenotypes can reveal insights into the evolution of unique traits. Here, we investigate a rare morphological innovation in modern haplochromine cichlids, a flap of fibrous tissue that causes a pronounced projection of the snout, which is limited to a single genus (Labeotropheus) of Lake Malawi cichlids. We compare flap size in our focal species L. fuelleborni (LF) to homologous landmarks in other closely related cichlid species that show a range of ecological overlap with LF, and demonstrate that variation in flap size is discontinuous among Malawi cichlid species. We demonstrate further that flap development in LF begins at early juvenile stages, and scales allometrically with body size. We then used an F2 hybrid mapping population, derived via crossing LF to a close ecological competitor that lacks this trait, Tropheops "red cheek" (TRC), to identify quantitative trait loci (QTL) that underlie flap development. In all, we identified four loci associated with variation in flap size, and for each the LF allele contributed to a larger flap. We next cross-referenced our QTL map with population genomic data, comparing natural populations of LF and TRC, to identify divergent polymorphisms within each QTL interval. Candidate genes for flap development are discussed. Together, these data indicate a relatively simple and tractable genetic basis for this morphological innovation, which is consistent with its apparently sudden and saltatory evolutionary history. J. Exp. Zool. (Mol. Dev. Evol.) 324B: 662-670, 2015. © 2015 Wiley Periodicals, Inc.
McCartney, T.; Scholz, C. A.; Shillington, D. J.; Accardo, N. J.; Chindandali, P. R. N.; Kamihanda, G.
Accommodation zones connect rift segments, which are fundamental elements of continental rift architecture. The sedimentary record aids our assessment of the evolution of this linkage. The central basin of Lake Malawi is one of the most structurally complex regions of the Malawi Rift. Border fault margins have been interpreted on both shorelines; three structures within the basin have been interpreted as segments of a corresponding accommodation zone. We investigate these structures by integrating single- and multi-channel reflection seismic data, including new MCS acquired in 2015 for the SEGMeNT project. The stratigraphic record in the central basin, inferred from seismic reflection profiles, provides compelling evidence that most fault-related subsidence is accommodated by the western border fault. Strata on both sides of all three structures dip to the west. The pre-rift basement in the sub-basin west of the central structure is considerably deeper (~ 4 s TWTT sub-bottom) than that in the broader eastern sub-basin (~ 2.5 s TWTT sub-bottom). A syncline in the eastern sub-basin shows little variation in seismic facies, particularly over the last 1.3 m.y. In contrast, the western sub-basin exhibits seismic facies indicative of fluvial input from two major rivers, siliciclastic input from the border fault footwall rising > 1000 m above lake level, and mud diapirs in the deepest part of the sub-basin. Horizons pierced by these diapirs onlap the central structure, suggesting diapir rise postdates relative uplift of the structure. Correlations with the age model from a 2005 scientific drilling project will better constrain this timing. The structural high helps focus siliciclastic sediments into the sub-basin, resulting in the overpressure conditions required for mud diapirism. We hypothesize that the diapirs are the result of sediment loading in the deep main depocenter of the central basin rather than fault mechanisms. The basement highs in the central basin control
Takeuchi, Yuichi; Hori, Michio; Tada, Shinya; Oda, Yoichi
The scale-eating cichlid Perissodus microlepis with asymmetric mouth is an attractive model of behavioral laterality: each adult tears off scales from prey fishes' left or right flanks according to the direction in which its mouth is skewed. To investigate the development of behavioral laterality and mouth asymmetry, we analyzed stomach contents and lower jaw-bone asymmetry of various-sized P. microlepis (22 ≤ SL<115 mm) sampled in Lake Tanganyika. The shapes of the pored scales found in each specimen's stomach indicated its attack side preference. Early-juvenile specimens (SL<45 mm) feeding mainly on zooplankton exhibited slight but significant mouth asymmetry. As the fish acquired scale-eating (45 mm ≤ SL), attack side preference was gradually strengthened, as was mouth asymmetry. Among size-matched individuals, those with more skewed mouths ate more scales. These findings show that behavioral laterality in scale-eating P. microlepis is established in association with development of mouth asymmetry which precedes the behavioral acquisition, and that this synergistic interaction between physical and behavioral literalities may contribute to efficient scale-eating.
Takeuchi, Yuichi; Hori, Michio; Tada, Shinya; Oda, Yoichi
The scale-eating cichlid Perissodus microlepis with asymmetric mouth is an attractive model of behavioral laterality: each adult tears off scales from prey fishes’ left or right flanks according to the direction in which its mouth is skewed. To investigate the development of behavioral laterality and mouth asymmetry, we analyzed stomach contents and lower jaw-bone asymmetry of various-sized P. microlepis (22≤SL<115mm) sampled in Lake Tanganyika. The shapes of the pored scales found in each specimen’s stomach indicated its attack side preference. Early-juvenile specimens (SL<45mm) feeding mainly on zooplankton exhibited slight but significant mouth asymmetry. As the fish acquired scale-eating (45mm≤SL), attack side preference was gradually strengthened, as was mouth asymmetry. Among size-matched individuals, those with more skewed mouths ate more scales. These findings show that behavioral laterality in scale-eating P. microlepis is established in association with development of mouth asymmetry which precedes the behavioral acquisition, and that this synergistic interaction between physical and behavioral literalities may contribute to efficient scale-eating. PMID:26808293
Ota, K; Kohda, M
Field and laboratory studies were conducted to examine how territorial males of a Lake Tanganyika cichlid Telmatochromis vittatus balance the conflicting demands on nest choice between occupying large nests with more females and avoiding reproductive parasitism (nest piracy, which is adopted by the largest males in the population). Pirates less frequently intruded the nests farther from neighbours, perhaps due to the costs associated with travelling between nests. The field experiment showed that territorial male T. vittatus sacrificed the fitness benefits that large nests offer and instead prioritized occupying the nests farther from neighbours on which fewer pirates intruded. The field observations suggested that they adopt different strategies for nest choice according to their relative competitive ability to pirates; the large territorial males, who are size-matched to pirates and can defend their nests against them, compete for larger nests among the more-isolated nests, whereas subordinate territorial males, which are smaller than pirates and thus inferior to them, compete for the more-isolated nests among the less-isolated nests. These findings suggest that the territorial male T. vittatus chooses the more-isolated nests to avoid pirate males at the expense of occupying large nests.
Castillo Cajas, Ruth F; Selz, Oliver M; Ripmeester, Erwin A P; Seehausen, Ole; Maan, Martine E
Environmental variation in signalling conditions affects animal communication traits, with possible consequences for sexual selection and reproductive isolation. Using spectrophotometry, we studied how male coloration within and between populations of two closely related Lake Victoria cichlid species (Pundamilia pundamilia and P. nyererei) covaries with water transparency. Focusing on coloration patches implicated in sexual selection, we predicted that in clear waters, with broad-spectrum light, (1) colours should become more saturated and (2) shift in hue away from the dominant ambient wavelengths, compared to more turbid waters. We found support for these predictions for the red and yellow coloration of P. nyererei but not the blue coloration of P. pundamilia. This may be explained by the species difference in depth distribution, which generates a steeper gradient in visual conditions for P. nyererei compared to P. pundamilia. Alternatively, the importance of male coloration in intraspecific sexual selection may differ between the species. We also found that anal fin spots, that is, the orange spots on male haplochromine anal fins that presumably mimic eggs, covaried with water transparency in a similar way for both species. This is in contrast to the other body regions studied and suggests that, while indeed functioning as signals, these spots may not play a role in species differentiation.
Miyagi, Ryutaro; Terai, Yohey; Aibara, Mitsuto; Sugawara, Tohru; Imai, Hiroo; Tachida, Hidenori; Mzighani, Semvua Isa; Okitsu, Takashi; Wada, Akimori; Okada, Norihiro
Reproductive isolation that prevents interspecific hybridization between closely related coexisting species maintains sympatric species diversity. One of the reproductive isolations is mate choice based on color signals (breeding color perceived by color vision). This is well known in several animal taxa, yet little is known about its genetic and molecular mechanism. Lake Victoria cichlid fishes are thought to be an example of sympatric species diversity. In the species inhabiting different light environments in rocky shore, speciation by sensory drive through color signals has been proposed by analyses of the long wavelength-sensitive (LWS) opsin gene and the male nuptial coloration. However, the genetic and molecular mechanism of how diversity of sympatric species occurring in the same habitat is maintained remains unknown. To address this issue, we determined nucleotide sequences of eight opsins of six sympatric species collected from a sandy-muddy shore--an ideal model system for studying sympatric species. Among eight opsins, the LWS and RH1 alleles were diversified and one particular allele is dominant or fixed in each species, and we propose that this is due to natural selection. The functions of their LWS alleles were also diversified as shown by absorption measurements of reconstituted visual pigments. To analyze the relationship between nuptial coloration and the absorption of LWS pigments, we systematically evaluated and defined nuptial coloration. We showed that the coloration was species specific with respect to hue and significantly differentiated by the index values of hue (dominant wavelength: λ(d)). The λ(d) value of the male nuptial coloration correlated with the absorption of LWS pigments from all the species, suggesting that reproductive isolation through mate choice using color signals may prevent sympatric interspecific hybridization, thereby maintaining the species diversity in sympatric species in Lake Victoria.
Johnson, Thomas; Werne, Josef
Hominin evolution underwent important changes in the last 1.3 million years, including the extinction of Paranthropus at about 1.2 Ma, leaving Homo as the sole hominin genus. Our genus experienced a major increase in cranial capacity at about 500 ka, and our species, H. sapiens, first appeared at ~200 ka. There was a major turnover in mammalian species in East Africa between 540 and 400 ka, favoring descendants of smaller size and less specialized diet. An understanding of what drove evolution in these directions is fundamental to understanding the development of modern H. sapiens. Climate certainly played a role, for it is the principal factor that influences the distribution of vegetation and habitability on the landscape. We present a 1.3 million year record of temperature and hydroclimate in the basin of Lake Malawi, the second deepest lake in Africa, derived from a 380 m sediment sequence taken from a water depth of 590 m by the Lake Malawi Drilling Project. Seismic reflection profiles used to select the site portray an undisturbed sedimentary section that was not impacted by erosion, turbidity currents or mass wasting events. Sediment samples were analyzed to produce records of temperature (TEX86) and aridity (Ca content and leaf wax δ13C). The temperature record displays progressively larger amplitude glacial-interglacial variations from MIS 13 (~500 ka) to MIS 5 (~125 ka). Intervals of low Ca abundance, which reflect lake high stands, correlate with times of depleted δ13Cwax and relatively warm temperatures. The Malawi basin experienced warm, wet interglacials and cooler (by about 2 - 4°C), dry glacial periods, with roughly a 100 ky periodicity since the Mid-Pleistocene Transition (MPT), about 900 ka. The paleoclimate record from Lake Malawi sediments portrays a transition from a highly variable and predominantly arid climate prior to 900 ka to a progressively more humid environment after the MPT dominated by 100 ky cycles consisting of warm, wet
Lane, Christine S.; Chorn, Ben T.; Johnson, Thomas C.
The most explosive volcanic event of the Quaternary was the eruption of Mt. Toba, Sumatra, 75,000 y ago, which produced voluminous ash deposits found across much of the Indian Ocean, Indian Peninsula, and South China Sea. A major climatic downturn observed within the Greenland ice cores has been attributed to the cooling effects of the ash and aerosols ejected during the eruption of the Youngest Toba Tuff (YTT). These events coincided roughly with a hypothesized human genetic bottleneck, when the number of our species in Africa may have been reduced to near extinction. Some have speculated that the demise of early modern humans at that time was due in part to a dramatic climate shift triggered by the supereruption. Others have argued that environmental conditions would not have been so severe to have such an impact on our ancestors, and furthermore, that modern humans may have already expanded beyond Africa by this time. We report an observation of the YTT in Africa, recovered as a cryptotephra layer in Lake Malawi sediments, >7,000 km west of the source volcano. The YTT isochron provides an accurate and precise age estimate for the Lake Malawi paleoclimate record, which revises the chronology of past climatic events in East Africa. The YTT in Lake Malawi is not accompanied by a major change in sediment composition or evidence for substantial temperature change, implying that the eruption did not significantly impact the climate of East Africa and was not the cause of a human genetic bottleneck at that time. PMID:23630269
Vanhove, Maarten P M; Snoeks, Jos; Volckaert, Filip A M; Huyse, Tine
Lake Tanganyika harbours the most diverse endemic cichlid fish assemblage of Africa, but its monogenean fish parasites have not been investigated. Here we report, for the first time, on the Gyrodactylus parasites in this hotspot of fish biodiversity. Haptor morphometrics and nuclear ribosomal DNA sequences revealed 3 new species on Zambian Simochromis diagramma: Gyrodactylus sturmbaueri n. sp., G. thysi n. sp. and G. zimbae n. sp. Their distinct morphology and strong genetic differentiation suggest that they belong to distant lineages within the genus Gyrodactylus, and phylogenetic reconstructions suggest affinities with other genera of gyrodactylids. Additional U-shaped haptoral plates in G. thysi n. sp. and a second large spine-like structure in the male copulatory organ of G. zimbae seem to represent new features for the genus. Such large diversity on a single host species can probably be explained by host-switching events during the course of evolution, in agreement with the generally accepted concept that ecological transfer is an important aspect of gyrodactylid speciation. Additional parasitological surveys on other host species, covering a broader phylogenetic and geographical range, should clarify the evolutionary history of Gyrodactylidae on cichlids in the African Great Lake and other parts of Africa.
Kmentová, Nikol; Gelnar, Milan; Mendlová, Monika; Van Steenberge, Maarten; Koblmüller, Stephan; Vanhove, Maarten P. M.
Lake Tanganyika is well-known for its high species-richness and rapid radiation processes. Its assemblage of cichlid fishes recently gained momentum as a framework to study parasite ecology and evolution. It offers a rare chance to investigate the influence of a deepwater lifestyle in a freshwater fish-parasite system. Our study represents the first investigation of parasite intraspecific genetic structure related to host specificity in the lake. It focused on the monogenean flatworm Cichlidogyrus casuarinus infecting deepwater cichlids belonging to Bathybates and Hemibates. Morphological examination of C. casuarinus had previously suggested a broad host range, while the lake’s other Cichlidogyrus species are usually host specific. However, ongoing speciation or cryptic diversity could not be excluded. To distinguish between these hypotheses, we analysed intraspecific diversity of C. casuarinus. Monogeneans from nearly all representatives of the host genera were examined using morphometrics, geomorphometrics and genetics. We confirmed the low host-specificity of C. casuarinus based on morphology and nuclear DNA. Yet, intraspecific variation of sclerotized structures was observed. Nevertheless, the highly variable mitochondrial DNA indicated recent population expansion, but no ongoing parasite speciation, confirming, for the first time in freshwater, reduced parasite host specificity in the deepwater realm, probably an adaptation to low host availability. PMID:28004766
Jackson, L. J.; Stone, J.; Cohen, A. S.
Debate over long, and short-term climatic impacts of the Mt. Toba super-eruption circa 75ky is often focused on East Africa. A severe drop in anatomically modern human populations has been hypothesized to be synchronous with a volcanic winter caused by the Toba super-eruption. If the Toba eruption caused a volcanic winter in East Africa, climatologically-sensitive ecosystems, such as Lake Malawi and its immediate watershed should show a direct and observable response in the sediment record. Cooler temperatures would cause a reduction of density contrast between epilimnion and hypolimnion waters, allowing for increased mixing and oxygenation of normally anoxic bottom waters. Enhanced mixing would cause noticeable changes in lake fly and algal communities. Cooler temperatures might also affect precipitation and the fire regime in the surrounding watershed. We analyzed two Lake Malawi cores at the finest practical resolution. Core 2A-10H-2 was analyzed in less than 6-year intervals and core 1C-8H-1 in 7-year intervals surrounding the Youngest Toba Tephra (YTT) for microfaunal abundance and variability, sediment composition, and evidence of changes in the occurrence of fires or watershed precipitation. Our analysis included point counts of diatoms and other algae, lake flies, charcoal, and siliciclastics. Changes in microfossil assemblage, variability, and abundance, as well as sediment composition around the YTT in Core 2A and 1C do not indicate that increased mixing or cooler temperatures occurred in either the central or northern basins of Lake Malawi. Similarly, charcoal counts do not suggest a change in fire regime. Our results indicate that at a subdecadal scale there was no substantial response in Lake Malawi or its immediate watershed to the Mt. Toba super-eruption, in contrast to predictions from the volcanic winter hypothesis.
Joost, Stéphane; Kalbermatten, Michael; Bezault, Etienne; Seehausen, Ole
When searching for loci possibly under selection in the genome, an alternative to population genetics theoretical models is to establish allele distribution models (ADM) for each locus to directly correlate allelic frequencies and environmental variables such as precipitation, temperature, or sun radiation. Such an approach implementing multiple logistic regression models in parallel was implemented within a computing program named MATSAM: . Recently, this application was improved in order to support qualitative environmental predictors as well as to permit the identification of associations between genomic variation and individual phenotypes, allowing the detection of loci involved in the genetic architecture of polymorphic characters. Here, we present the corresponding methodological developments and compare the results produced by software implementing population genetics theoretical models (DFDIST: and BAYESCAN: ) and ADM (MATSAM: ) in an empirical context to detect signatures of genomic divergence associated with speciation in Lake Victoria cichlid fishes.
Scholz, C. A.; Wood, D. A.; Shillington, D. J.; McCartney, T.; Accardo, N. J.
The western branch of the East African Rift is characterized by modest amounts of mainly amagmatic extension; deeply-subsided, fault-controlled basins; and large-magnitude, deep seismicity. Lakes Tanganyika and Malawi are two of the world's largest lakes, with maximum water depths of 1450 and 700 m respectively. Newly acquired seismic reflection data, along with newly reprocessed legacy data reveal thick sedimentary sections, in excess of 5 km in some localities. The 1980's vintage legacy data from Project PROBE have been reprocessed through pre-stack depth migration in Lake Tanganyika, and similar reprocessing of legacy data from Lake Malawi is forthcoming. New high-fold and large-source commercial and academic data have recently been collected in southern Lake Tanganyika, and in the northern and central basins of Lake Malawi as part of the 2015 SEGMeNT project. In the case of Lake Tanganyika, new data indicate the presence of older sediment packages that underlie previously identified "pre-rift" basement (the "Nyanja Event"). These episodes of sedimentation and extension may substantially predate the modern lake. These deep stratal reflections are absent in many localites, possibly on account of attenuation of the acoustic signal. However in one area of southern Lake Tanganyika, the newly-observed deep strata extend axially for ~70 km, likely representing deposits from a discrete paleolake. The high-amplitude Nyanja Event is interpreted as the onset of late-Cenozoic rifting, and the changing character of the overlying depositional sequences reflects increasing relief in the rift valley, as well as the variability of fluvial inputs, and the intermittent connectivity of upstream lake catchments. Earlier Tanganyika sequences are dominated by shallow lake and fluvial-lacustrine facies, whereas later sequences are characterized by extensive gravity flow deposition in deep water, and pronounced erosion and incision in shallow water depths and on littoral platforms. The
Brown, E. T.; Johnson, T. C.; Scholz, C. A.; King, J.; Cohen, A. S.
As part of the Lake Malawi Drilling Project we have undertaken a 1 cm-scale resolution XRF scanning study of core from site 2A in the Northern Basin of the lake (ca. 10 South latitude). This 38-m core provides an essentially continuous record of regional climate over the past 70,000 years, so our XRF analyses correspond to an average temporal resolution of approximately 20 years. We have focused our initial study on certain parameters, including Fe, Zr:Ti, and Si:Ti. These may be interpreted to represent changing input of terrigenous sediments (delivered to the lake rivers), soils developed from weathered volcanic ash (delivered more effectively under drier windier conditions), and biogenic silica, respectively. Our initial analyses of the XRF results indicate abrupt and strong, millennial-scale variability in regional climate throughout the past 70,000 years. The series of events consists of rapid (~100-year) increases in Zr:Ti and Si:Ti followed by slower decreases toward background values. These events are strikingly similar in form and in timing to Dansgaard- Oeschger events of high northern latitudes. The Younger Dryas and the Holocene "cold" periods in the Northern Hemisphere were times of enhanced input of wind-blown volcanic ash-derived materials and of diatom productivity, probably stimulated by northerly winds and upwelling in the north of the lake. Interestingly, in contrast to the Late Glacial and Holocene, times of higher productivity and enhanced input of volcanic-ash derived material in the earlier part of the record coincide with episodes of warming as recorded in Greenland ice. This suggests that a significant change in the tropical response to climate forcing occurred at the time of waning of Northern Hemisphere glaciation.
Martin, Christopher H
Models predict that sympatric speciation depends on restrictive parameter ranges, such as sufficiently strong disruptive selection and assortative mating, but compelling examples in nature have rarely been used to test these predictions. I measured the strength of assortative mating within a species complex of Tilapia in Lake Ejagham, Cameroon, a celebrated example of incipient sympatric adaptive radiation. This species complex is in the earliest stages of speciation: morphological and ecological divergence are incomplete, species differ primarily in breeding coloration, and introgression is common. I captured 27 mated pairs in situ and measured the diet, color, size, and morphology of each individual. I found strong assortative mating by color, size, head depth, and dietary source of benthic or pelagic prey along two independent dimensions of assortment. Thus, Ejagham Tilapia showed strong assortative mating most conducive to sympatric speciation. Nonetheless, in contrast to a morphologically bimodal Sarotherodon cichlid species pair in the lake, Ejagham Tilapia show more limited progress toward speciation, likely due to insufficient strength of disruptive selection on morphology estimated in a previous study (γ = 0.16). This supports the predicted dependence of sympatric speciation on strong assortment and strong disruptive selection by examining a potentially stalled example in nature.
Wilson, Laura A. B.; Colombo, Marco; Sánchez-Villagra, Marcelo R.; Salzburger, Walter
Phenotype-environment correlations and the evolution of trait interactions in adaptive radiations have been widely studied to gain insight into the dynamics underpinning rapid species diversification. In this study we explore the phenotype-environment correlation and evolution of operculum shape in cichlid fishes using an outline-based geometric morphometric approach combined with stable isotope indicators of macrohabitat and trophic niche. We then apply our method to a sample of extinct saurichthyid fishes, a highly diverse and near globally distributed group of actinopterygians occurring throughout the Triassic, to assess the utility of extant data to inform our understanding of ecomorphological evolution in extinct species flocks. A series of comparative methods were used to analyze shape data for 54 extant species of cichlids (N = 416), and 6 extinct species of saurichthyids (N = 44). Results provide evidence for a relationship between operculum shape and feeding ecology, a concentration in shape evolution towards present along with evidence for convergence in form, and significant correlation between the major axes of shape change and measures of gut length and body elongation. The operculum is one of few features that can be compared in extant and extinct groups, enabling reconstruction of phenotype-environment interactions and modes of evolutionary diversification in deep time. PMID:26584885
Hu, Yinan; Albertson, R. Craig
Adaptive variation in the craniofacial skeleton is a key component of resource specialization and habitat divergence in vertebrates, but the proximate genetic mechanisms that underlie complex patterns of craniofacial variation are largely unknown. Here we demonstrate that the Hedgehog (Hh) signaling pathway mediates widespread variation across a complex functional system that affects the kinematics of lower jaw depression—the opercular four-bar linkage apparatus—among Lake Malawi cichlids. By using a combined quantitative trait locus mapping and population genetics approach, we show that allelic variation in the Hh receptor, ptch1, affects the development of distinct bony elements in the head that represent two of three movable links in this functional system. The evolutionarily derived allele is found in species that feed from the water column, and is associated with shifts in anatomy that translate to a four-bar system capable of faster jaw rotation. Alternatively, the ancestral allele is found in species that feed on attached algae, and is associated with the development of a four-bar system that predicts slower jaw movement. Experimental manipulation of the Hh pathway during cichlid development recapitulates functionally salient natural variation in craniofacial geometry. In all, these results significantly extend our understanding of the mechanisms that fine-tune the craniofacial skeletal complex during adaptation to new foraging niches. PMID:24912175
Hu, Yinan; Albertson, R Craig
Adaptive variation in the craniofacial skeleton is a key component of resource specialization and habitat divergence in vertebrates, but the proximate genetic mechanisms that underlie complex patterns of craniofacial variation are largely unknown. Here we demonstrate that the Hedgehog (Hh) signaling pathway mediates widespread variation across a complex functional system that affects the kinematics of lower jaw depression--the opercular four-bar linkage apparatus--among Lake Malawi cichlids. By using a combined quantitative trait locus mapping and population genetics approach, we show that allelic variation in the Hh receptor, ptch1, affects the development of distinct bony elements in the head that represent two of three movable links in this functional system. The evolutionarily derived allele is found in species that feed from the water column, and is associated with shifts in anatomy that translate to a four-bar system capable of faster jaw rotation. Alternatively, the ancestral allele is found in species that feed on attached algae, and is associated with the development of a four-bar system that predicts slower jaw movement. Experimental manipulation of the Hh pathway during cichlid development recapitulates functionally salient natural variation in craniofacial geometry. In all, these results significantly extend our understanding of the mechanisms that fine-tune the craniofacial skeletal complex during adaptation to new foraging niches.
Parsons, Kevin J; Concannon, Moira; Navon, Dina; Wang, Jason; Ea, Ilene; Groveas, Kiran; Campbell, Calum; Albertson, R Craig
Phenotypic plasticity allows organisms to change their phenotype in response to shifts in the environment. While a central topic in current discussions of evolutionary potential, a comprehensive understanding of the genetic underpinnings of plasticity is lacking in systems undergoing adaptive diversification. Here, we investigate the genetic basis of phenotypic plasticity in a textbook adaptive radiation, Lake Malawi cichlid fishes. Specifically, we crossed two divergent species to generate an F3 hybrid mapping population. At early juvenile stages, hybrid families were split and reared in alternate foraging environments that mimicked benthic/scraping or limnetic/sucking modes of feeding. These alternate treatments produced a variation in morphology that was broadly similar to the major axis of divergence among Malawi cichlids, providing support for the flexible stem theory of adaptive radiation. Next, we found that the genetic architecture of several morphological traits was highly sensitive to the environment. In particular, of 22 significant quantitative trait loci (QTL), only one was shared between the environments. In addition, we identified QTL acting across environments with alternate alleles being differentially sensitive to the environment. Thus, our data suggest that while plasticity is largely determined by loci specific to a given environment, it may also be influenced by loci operating across environments. Finally, our mapping data provide evidence for the evolution of plasticity via genetic assimilation at an important regulatory locus, ptch1. In all, our data address long-standing discussions about the genetic basis and evolution of plasticity. They also underscore the importance of the environment in affecting developmental outcomes, genetic architectures, morphological diversity and evolutionary potential.
Yost, C. L.; Cohen, A. S.
The recent discovery of cryptotephra visually and chemically matched to the Youngest Toba Tuff (YTT, 75.0 ± 0.9 kyr) in Lake Malawi drill core sediments has spurred renewed interest in this period of time in East Africa. The YTT is the most recent and largest of the four Mount Toba eruptions, and is the only super-eruption to have taken place during the Quaternary. The timing of the YTT approximately coincides with a hypothesized human genetic bottleneck. Several climate models have proposed an episode of global cooling following the YTT; however, the magnitude and duration of the cooling is much debated, ranging from just a few degrees of cooling to a state of volcanic winter. Cored sediments from Lake Malawi provide an excellent record of local variability in the lake's watershed that may be linked to specific climatic events. To investigate the possible effects of the YTT in East Africa, we continuously sampled Lake Malawi drill core 2A-10H-2 at 2-4 mm (~6 yr) intervals above and below the first occurrence of the YTT. Poaceae phytoliths were grouped into plant functional types (C3, C4, xerophytic, mesophytic, arboreal, etc.), revealing mostly subtle changes in terrestrial vegetation over the ~400 yr time period examined. Abrupt increases in concentration values for phytoliths derived from riverine Podostemaceae plants appear to signal increased discharge from rivers draining the surrounding uplands. Perhaps most significant is the increasing trend in burned phytoliths and decreasing trend in tree phytoliths post-YTT. Although there appears to be a very weak cooling signal synchronous with the YTT, the most abrupt terrestrial vegetation changes appear to be better correlated with the deposition of a slightly older cryptotephra horizon derived from the local Rungwe Volcanic Province. A potential complication with this record is the existence of a turbidite pre-YTT that encompasses the Rungwe horizon.
Tsuboi, Masahito; Husby, Arild; Kotrschal, Alexander; Hayward, Alexander; Buechel, Séverine D; Zidar, Josefina; Løvlie, Hanne; Kolm, Niclas
The brain is one of the most energetically expensive organs in the vertebrate body. Consequently, the energetic requirements of encephalization are suggested to impose considerable constraints on brain size evolution. Three main hypotheses concerning how energetic constraints might affect brain evolution predict covariation between brain investment and (1) investment into other costly tissues, (2) overall metabolic rate, and (3) reproductive investment. To date, these hypotheses have mainly been tested in homeothermic animals and the existing data are inconclusive. However, there are good reasons to believe that energetic limitations might play a role in large-scale patterns of brain size evolution also in ectothermic vertebrates. Here, we test these hypotheses in a group of ectothermic vertebrates, the Lake Tanganyika cichlid fishes. After controlling for the effect of shared ancestry and confounding ecological variables, we find a negative association between brain size and gut size. Furthermore, we find that the evolution of a larger brain is accompanied by increased reproductive investment into egg size and parental care. Our results indicate that the energetic costs of encephalization may be an important general factor involved in the evolution of brain size also in ectothermic vertebrates.
Parente, Thiago E M; De-Oliveira, Ana C A X; Paumgartten, Francisco J R
The induction of cytochrome P4501A-mediated activity (e.g. ethoxyresorufin-O-deethylation, EROD) has been used as a biomarker for monitoring fish exposure to AhR-receptor ligands such as polycyclic aromatic hydrocarbons (PAH), polychlorinated biphenyls (PCB) and polychlorinated dibenzo-dioxins/furans (PCDD/Fs). In this study we found that hepatic EROD is induced in fish ("Nile tilapia", Oreochromis niloticus and "acará", Geophagus brasiliensis) from the Guandu River (7-17-fold) and Jacarepaguá Lake (7-fold), Rio de Janeiro, Brazil. Since both cichlid fish are consumed by the local population and the Guandu River is the main source of the drinking water supply for the greater Rio de Janeiro metropolitan area, pollution by cytochrome P4501A-inducing chemicals is a cause for concern and should be further investigated in sediments, water and biota. We additionally showed that EROD activity in the fish liver post-mitochondrial supernatant-simpler, cheaper and less time consuming to prepare than the microsomal fraction-is sufficiently sensitive for monitoring purposes.
Nagoli, Joseph; Chiwona-Karltun, Linley
This paper provides an in-depth understanding of social dynamics in the form of kinship ties in matrilineal societies. It unpacks gender roles and relationships at the community level to understand how social structures, created by the pattern of relations, enhance or hinder coping initiatives during lake recessions in the Lake Chilwa socio-ecological system. Using qualitative and quantitative methods including in-depth interviews, Focus Group Discussions (FGD) and household questionnaires, this paper provides insights into how people in the Lake Chilwa social-ecological system prepare for and cope with Lake Chilwa water recessions by capitalising on their social networks. The findings have shown that during lake recessions poor households in the Lake Chilwa socio-ecological system are cushioned by fellow households through lineage networks. There is strong tradition secured through kinship ties where the generic term 'mwambo wathu' (our tradition) embraces a group of formative norms enforced via a series of rules and rituals. Based in a matrilineal system, women have strong rights over land allocation and use in which the female sorority group 'mbumba' is very stable. There is a great deal of reciprocity and food redistribution among the households. This self-organization provides deeper meaning about how people assimilate the dynamics affecting their social and natural environments. Specifically the findings provide evidence on how households, communities and their livelihoods in Africa are reconstructed as natural resources fluctuate.
Kafumbata, Dalitso; Jamu, Daniel; Chiotha, Sosten
This paper reviews the importance of African lakes and their management challenges. African inland lakes contribute significantly to food security, livelihoods and national economies through direct exploitation of fisheries, water resources for irrigation and hydropower generation. Because of these key contributions, the ecosystem services provided are under significant stress mainly owing to high demand by increasing populations, negative anthropogenic impacts on lake catchments and high levels of poverty which result in unsustainable use. Climate variability exacerbates the stress on these ecosystems. Current research findings show that the lakes cannot sustain further development activities on the scale seen over the past few decades. Millions of people are at risk of losing livelihoods through impacts on livestock and wildlife. The review further shows that the problems facing these lakes are beyond the purview of current management practices. A much better understanding of the interactions and feedbacks between different components of the lake socio-ecological systems is needed to address the complex challenges of managing these ecosystem services. This review suggests that the three small wetlands of Chad, Chilwa and Naivasha provide an opportunity for testing novel ideas that integrate sustainability of natural resource management with livelihoods in order to inform policy on how future land use and climatic variability will affect both food security and the ecosystem services associated with it. PMID:24535395
Torres-Dowdall, Julián; Henning, Frederico; Elmer, Kathryn R; Meyer, Axel
The visual system in the colorful cichlid fishes from the African great lakes is believed to be important for their adaptive radiations. However, few studies have attempted to compare the visual system of radiating cichlid lineages with that of cichlids that have not undergone recent radiations. One such study published in this journal (Schott RK, Refvik SP, Hauser FE, López-Fernández H, Chang BSW. 2014. Divergent positive selection in rhodopsin from lake and riverine cichlid fishes. Mol Biol Evol. 31:1149-1165) found divergent selection on rhodopsin between African lacustrine and riverine cichlid species and riverine Neotropical cichlids, concluding that ecology drives the molecular evolution of this opsin. Here, we expand this analysis by incorporating rhodopsin sequences from Neotropical lacustrine cichlids and show that both ecology and phylogeny are important drivers of the molecular evolution of rhodopsin in cichlids. We found little overlap of sites under selection between African and Neotropical lineages and a faster rate of molecular evolution in African compared with Neotropical cichlids. These results support the notion that genetic or population genetic features particular to African cichlids contributed to their radiations.
Ford, Antonia G. P.; Rüber, Lukas; Newton, Jason; Dasmahapatra, Kanchon K.; Balarin, John D.; Bruun, Kristoffer; Day, Julia J.
Ecomorphological differentiation is a key feature of adaptive radiations, with a general trend for specialization and niche expansion following divergence. Ecological opportunity afforded by invasion of a new habitat is thought to act as an ecological release, facilitating divergence, and speciation. Here, we investigate trophic adaptive morphology and ecology of an endemic clade of oreochromine cichlid fishes (Alcolapia) that radiated along a herbivorous trophic axis following colonization of an isolated lacustrine environment, and demonstrate phenotype‐environment correlation. Ecological and morphological divergence of the Alcolapia species flock are examined in a phylogenomic context, to infer ecological niche occupation within the radiation. Species divergence is observed in both ecology and morphology, supporting the importance of ecological speciation within the radiation. Comparison with an outgroup taxon reveals large‐scale ecomorphological divergence but shallow genomic differentiation within the Alcolapia adaptive radiation. Ancestral morphological reconstruction suggests lake colonization by a generalist oreochromine phenotype that diverged in Lake Natron to varied herbivorous morphologies akin to specialist herbivores in Lakes Tanganyika and Malawi. PMID:27659769
Lund, Steve; Platzman, Ellen; Johnson, Tom
We have conducted a paleomagnetic study of Late Quaternary sediments from Lake Malawi, East Africa, in order to develop a high-resolution record of paleomagnetic secular variation (PSV). This study has recovered PSV records from two cores (3P, 6P) in northern Lake Malawi (10.0°S, 34.3°E). The PSV appears to be recorded in fine-grained detrital magnetite/titanomagnetite grains. Detailed af demagnetization of the natural remanence (NRM) shows that a distinctive characteristic remanence (ChRM) is demagnetized from ∼20 to 80 mT, which decreases simply toward the origin. The resulting directional PSV records for 3P and 6P are easily correlatable with 29 distinct inclination features and 29 declination features. The statistical character of the PSV in both cores is consistent with Holocene PSV noted at other Holocene equatorial sites. Radiocarbon dating of the cores is based on 18 independent radiocarbon dates and four dated stratigraphic horizons that can be correlated into each core. The final directional PSV time series cover the last 24,000 years with an average sediment accumulation rate of ∼30 cm/kyr. We have also developed a relative paleointensity estimate for these PSV records based on normalizing the NRM (after 20 mT af demagnetization) by the SIRM (after 20 mT af demagnetization). Changing sedimentation patterns complicate any attempt to develop a single paleointensity record for the entire core lengths. We have developed a relative paleointensity record for the last 6000 years that has 14 correlatable features including 5 notable peaks in intensity. Three of these peaks are synchronous with paleointensity highs farther north in SE Europe/SW Asia/Egypt but two of the peaks are at times of low paleointensity farther north. We interpret this to indicate that Lake Malawi (10°S) is at least partly under the influence of a different flux-regeneration region of the outer-core dynamo. A relative paleointensity record was also developed for ∼11,000-24,000 YBP
Maan, Martine E; Hofker, Kees D; van Alphen, Jacques J M; Seehausen, Ole
The role of selection in speciation is a central yet poorly understood problem in evolutionary biology. The rapid radiations of extremely colorful cichlid fish in African lakes have fueled the hypothesis that sexual selection can drive species divergence without geographical isolation. Here we present experimental evidence for a mechanism by which sexual selection becomes divergent: in two sibling species from Lake Victoria, female mating preferences for red and blue male nuptial coloration coincide with their context-independent sensitivities to red and blue light, which in turn correspond to a difference in ambient light in the natural habitat of the species. These results suggest that natural selection on visual performance, favoring different visual properties in different spectral environments, may lead to divergent sexual selection on male nuptial coloration. This interplay of ecological and sexual selection along a light gradient may provide a mechanism of rapid speciation through divergent sensory drive.
Marshall, N. Justin; Cronin, Thomas W.; Seehausen, Ole; Carleton, Karen L.
A major goal of evolutionary biology is to unravel the molecular genetic mechanisms that underlie functional diversification and adaptation. We investigated how changes in gene regulation and coding sequence contribute to sensory diversification in two replicate radiations of cichlid fishes. In the clear waters of Lake Malawi, differential opsin expression generates diverse visual systems, with sensitivities extending from the ultraviolet to the red regions of the spectrum. These sensitivities fall into three distinct clusters and are correlated with foraging habits. In the turbid waters of Lake Victoria, visual sensitivity is constrained to longer wavelengths, and opsin expression is correlated with ambient light. In addition to regulatory changes, we found that the opsins coding for the shortest- and longest-wavelength visual pigments have elevated numbers of potentially functional substitutions. Thus, we present a model of sensory evolution in which both molecular genetic mechanisms work in concert. Changes in gene expression generate large shifts in visual pigment sensitivity across the collective opsin spectral range, but changes in coding sequence appear to fine-tune visual pigment sensitivity at the short- and long-wavelength ends of this range, where differential opsin expression can no longer extend visual pigment sensitivity. PMID:20027211
Hofmann, Christopher M; O'Quin, Kelly E; Marshall, N Justin; Cronin, Thomas W; Seehausen, Ole; Carleton, Karen L
A major goal of evolutionary biology is to unravel the molecular genetic mechanisms that underlie functional diversification and adaptation. We investigated how changes in gene regulation and coding sequence contribute to sensory diversification in two replicate radiations of cichlid fishes. In the clear waters of Lake Malawi, differential opsin expression generates diverse visual systems, with sensitivities extending from the ultraviolet to the red regions of the spectrum. These sensitivities fall into three distinct clusters and are correlated with foraging habits. In the turbid waters of Lake Victoria, visual sensitivity is constrained to longer wavelengths, and opsin expression is correlated with ambient light. In addition to regulatory changes, we found that the opsins coding for the shortest- and longest-wavelength visual pigments have elevated numbers of potentially functional substitutions. Thus, we present a model of sensory evolution in which both molecular genetic mechanisms work in concert. Changes in gene expression generate large shifts in visual pigment sensitivity across the collective opsin spectral range, but changes in coding sequence appear to fine-tune visual pigment sensitivity at the short- and long-wavelength ends of this range, where differential opsin expression can no longer extend visual pigment sensitivity.
Terai, Yohey; Mayer, Werner E.; Klein, Jan; Tichy, Herbert; Okada, Norihiro
In East African Lake Victoria >200 endemic species of haplochromine fishes have been described on the basis of morphological and behavioral differences. Yet molecular analysis has failed to reveal any species-specific differences among these fishes in either mitochondrial or nuclear genes. Although the genes could be shown to vary, the variations represent trans-species polymorphisms not yet assorted along species lines. Nevertheless, fixed genetic differences must exist between the species at loci responsible for the adaptive characters distinguishing the various forms from one another. Here we describe variation and fixation at the long wavelength-sensitive (LWS) opsin locus, which is selection-driven, adaptive, and if not species- then at least population-specific. Because color is one of the characters distinguishing species of haplochromine fishes and color perception plays an important part in food acquisition and mate choice, we suggest that the observed variation and fixation at the LWS opsin locus may have been involved in the process that has led to the spectacular species divergence of haplochromine fishes in Lake Victoria. PMID:12438648
Magalhaes, I S; Smith, A M; Joyce, D A
The strategies and traits males evolve to mate with females are incredible in their diversity. Theory on the evolution of secondary sexual characters suggests that evolving any costly trait or strategy will pay off and stabilise in the population if it is advantageous compared to the alternative less costly strategy, but quantifying the relative success of the two can be difficult. In Lake Malawi, Africa, there are >200 species of cichlid fish in which the males form leks and spend several weeks per year building sand-castle "bowers" several times their size. We tested the idea that a less costly "sneaking" strategy could be successful by quantifying the mating success of bower-holding versus non-bower-holding males. We PIT-tagged every fish in a semi-natural experimental set-up and placed tag-readers on the side of bowers to determine which fish held a bower. We then genotyped the eggs removed from females' mouths to assign paternity of each egg. Broods were fathered by up to 3 different males. Although paternity was mostly assigned to males that held a bower, a small number of males who did not own a bower were more successful than some of those that did, indicating a role for an alternative strategy in these bower builders.
Magalhaes, I. S.; Smith, A. M.; Joyce, D. A.
The strategies and traits males evolve to mate with females are incredible in their diversity. Theory on the evolution of secondary sexual characters suggests that evolving any costly trait or strategy will pay off and stabilise in the population if it is advantageous compared to the alternative less costly strategy, but quantifying the relative success of the two can be difficult. In Lake Malawi, Africa, there are >200 species of cichlid fish in which the males form leks and spend several weeks per year building sand-castle “bowers” several times their size. We tested the idea that a less costly “sneaking” strategy could be successful by quantifying the mating success of bower-holding versus non-bower-holding males. We PIT-tagged every fish in a semi-natural experimental set-up and placed tag-readers on the side of bowers to determine which fish held a bower. We then genotyped the eggs removed from females’ mouths to assign paternity of each egg. Broods were fathered by up to 3 different males. Although paternity was mostly assigned to males that held a bower, a small number of males who did not own a bower were more successful than some of those that did, indicating a role for an alternative strategy in these bower builders. PMID:28128313
Henning, Frederico; Lee, Hyuk Je; Franchini, Paolo; Meyer, Axel
The genetic dissection of naturally occurring phenotypes sheds light on many fundamental and longstanding questions in speciation and adaptation and is a central research topic in evolutionary biology. Until recently, forward-genetic approaches were virtually impossible to apply to nonmodel organisms, but the development of next-generation sequencing techniques eases this difficulty. Here, we use the ddRAD-seq method to map a colour trait with a known adaptive function in cichlid fishes, well-known textbook examples for rapid rates of speciation and astonishing phenotypic diversification. A suite of phenotypic key innovations is related to speciation and adaptation in cichlids, among which body coloration features prominently. The focal trait of this study, horizontal stripes, evolved in parallel in several cichlid radiations and is associated with piscivorous foraging behaviour. We conducted interspecific crosses between Haplochromis sauvagei and H. nyererei and constructed a linkage map with 867 SNP markers distributed on 22 linkage groups and total size of 1130.63 cM. Lateral stripes are inherited as a Mendelian trait and map to a single genomic interval that harbours a paralog of a gene with known function in stripe patterning. Dorsolateral and mid-lateral stripes were always coinherited and are thus under the same genetic control. Additionally, we directly quantify the genotyping error rates in RAD markers and offer guidelines for identifying and dealing with errors. Uncritical marker selection was found to severely impact linkage map construction. Fortunately, by applying appropriate quality control steps, a genotyping accuracy of >99.9% can be reached, thus allowing for efficient linkage mapping of evolutionarily relevant traits.
Schwalbe, Margot A B; Sevey, Benjamin J; Webb, Jacqueline F
The mechanosensory lateral line system of fishes detects water motions within a few body lengths of the source. Several types of artificial stimuli have been used to probe lateral line function in the laboratory, but few studies have investigated the role of flow sensing in benthic feeding teleosts. In this study, we used artificial flows emerging from a sandy substrate to assess the contribution of flow sensing to prey detection in the peacock cichlid, Aulonocara stuartgranti, which feeds on benthic invertebrates in Lake Malawi. Using a positive reinforcement protocol, we trained fish to respond to flows lacking the visual and chemical cues generated by tethered prey in prior studies with A. stuartgranti Fish successfully responded to artificial flows at all five rates presented (characterized using digital particle image velocimetry), and showed a range of flow-sensing behaviors, including an unconditioned bite response. Immediately after lateral line inactivation, fish rarely responded to flows and the loss of vital fluorescent staining of hair cells (with 4-di-2-ASP) verified lateral line inactivation. Within 2 days post-treatment, some aspects of flow-sensing behavior returned and after 7 days, flow-sensing behavior and hair cell fluorescence both returned to pre-treatment levels, which is consistent with the reported timing of hair cell regeneration in other vertebrates. The presentation of ecologically relevant water flows to assess flow-sensing behaviors and the use of a positive reinforcement protocol are methods that present new opportunities to study the role of flow sensing in the feeding ecology of benthic feeding fishes.
Fischer, Christoph; Koblmüller, Stephan; Gülly, Christian; Schlötterer, Christian; Sturmbauer, Christian; Thallinger, Gerhard G.
The cichlid fishes of the East African Great Lakes represent a model especially suited to study adaptive radiation and speciation. With several African cichlid genome projects being in progress, a promising set of closely related genomes is emerging, which is expected to serve as a valuable data base to solve questions on genotype-phenotype relations. The mitochondrial (mt) genomes presented here are the first results of the assembly and annotation process for two closely related but eco-morphologically highly distinct Lake Tanganyika cichlids, Petrochromis trewavasae and Tropheus moorii. The genomic sequences comprise 16,588 bp (P. trewavasae) and 16,590 bp (T. moorii), and exhibit the typical mitochondrial structure, with 13 protein-coding genes, 2 rRNA genes, 22 tRNA genes, and a non-coding control region. Analyses confirmed that the two species are very closely related with an overall sequence similarity of 96%. We analyzed the newly generated sequences in the phylogenetic context of 21 published labroid fish mitochondrial genomes. Consistent with other vertebrates, the D-loop region was found to evolve faster than protein-coding genes, which in turn are followed by the rRNAs; the tRNAs vary greatly in the rate of sequence evolution, but on average evolve the slowest. Within the group of coding genes, ND6 evolves most rapidly. Codon usage is similar among examined cichlid tribes and labroid families; although a slight shift in usage patterns down the gene tree could be observed. Despite having a clearly different nucleotide composition, ND6 showed a similar codon usage. C-terminal ends of Cox1 exhibit variations, where the varying number of amino acids is related to the structure of the obtained phylogenetic tree. This variation may be of functional relevance for Cox1 synthesis. PMID:23826193
Machado-Schiaffino, Gonzalo; Kautt, Andreas F; Torres-Dowdall, Julian; Baumgarten, Lukas; Henning, Frederico; Meyer, Axel
Sympatric speciation has been debated in evolutionary biology for decades. Although it has gained in acceptance recently, still only a handful of empirical examples are seen as valid (e.g. crater lake cichlids). In this study, we disentangle the role of hypertrophied lips in the repeated adaptive radiations of Nicaraguan crater lake cichlid fish. We assessed the role of disruptive selection and assortative mating during the early stages of divergence and found a functional trade-off in feeding behaviour between thick- and thin-lipped ecotypes, suggesting that this trait is a target of disruptive selection. Thick-lipped fish perform better on nonevasive prey at the cost of a poorer performance on evasive prey. Using enclosures in the wild, we found that thick-lipped fish perform significantly better in rocky than in sandy habitats. We found almost no mixed pairs during two breeding seasons and hence significant assortative mating. Genetic differentiation between ecotypes seems to be related to the time since colonization, being subtle in L. Masaya (1600 generations ago) and absent in the younger L. Apoyeque (<600 generations ago). Genome-wide differentiation between ecotypes was higher in the old source lakes than in the young crater lakes. Our results suggest that hypertrophied lips might be promoting incipient sympatric speciation through divergent selection (ecological divergence in feeding performance) and nonrandom mating (assortative mating) in the young Nicaraguan crater lakes. Nonetheless, further manipulative experiments are needed in order to confirm the role of hypertrophied lips as the main cue for assortative mating.
Nikaido, Masato; Ota, Tomoki; Hirata, Tadashi; Suzuki, Hikoyu; Satta, Yoko; Aibara, Mitsuto; Mzighani, Semvua I; Sturmbauer, Christian; Hagino-Yamagishi, Kimiko; Okada, Norihiro
Fish use olfaction to detect a variety of nonvolatile chemical signals, and thus, this sense is key to survival and communication. However, the contribution of the olfactory sense to social-especially reproductive-interactions in cichlids is still controversial. To obtain insights into this issue, we investigated the genes encoding V1Rs-possible candidates for reproductive pheromone receptors-among East-African cichlids. Interestingly, we found an excess of nonsynonymous over synonymous substitutions in four of six V1R genes in multiple cichlid lineages. First, we found that highly dimorphic V1R2 allele groups were shared among the cichlids inhabiting all East-African Great Lakes emerged through the episodic accumulation of the nonsynonymous substitutions prior to the radiation of the Lake Tanganyika species flock. We further detected such episodic events in V1R1 of the tribe Tropheini, and in V1R3 and V1R6 of the tribe Trematocarini. The excess of nonsynonymous substitutions in these examples were indicated as dN/dS > 1, which were all statistically significant by Fisher's exact test. Furthermore, we speculate that the amino acid changes in these episodic events are likely functional switch because they occurred in the putative ligand-binding pocket. Our finding of the occurrence of multiple episodic events and the unexpected gene diversity in one unique gene family is suggestive of the contribution of the V1R to the species diversification and the social interaction in cichlids.
Franchini, Paolo; Xiong, Peiwen; Fruciano, Carmelo; Meyer, Axel
Cichlid fishes are an ideal model system for studying biological diversification because they provide textbook examples of rapid speciation. To date, there has been little focus on the role of gene regulation during cichlid speciation. However, in recent years, gene regulation has been recognized as a powerful force linking diversification in gene function to speciation. Here, we investigated the potential role of miRNA regulation in the diversification of six cichlid species of the Midas cichlid lineage (Amphilophus spp.) inhabiting the Nicaraguan crater lakes. Using several genomic resources, we inferred 236 Midas miRNA genes that were used to predict the miRNA target sites on 8,232 Midas 3′-UTRs. Using population genomic calculations of SNP diversity, we found the miRNA genes to be more conserved than protein coding genes. In contrast to what has been observed in other cichlid fish, but similar to what has been typically found in other groups, we observed genomic signatures of purifying selection on the miRNA targets by comparing these sites with the less conserved nontarget portion of the 3′-UTRs. However, in one species pair that has putatively speciated sympatrically in crater Lake Apoyo, we recovered a different pattern of relaxed purifying selection and high genetic divergence at miRNA targets. Our results suggest that sequence evolution at miRNA binding sites could be a critical genomic mechanism contributing to the rapid phenotypic evolution of Midas cichlids. PMID:27189980
Background Phenotypic evolution and its role in the diversification of organisms is a central topic in evolutionary biology. A neglected factor during the modern evolutionary synthesis, adaptive phenotypic plasticity, more recently attracted the attention of many evolutionary biologists and is now recognized as an important ingredient in both population persistence and diversification. The traits and directions in which an ancestral source population displays phenotypic plasticity might partly determine the trajectories in morphospace, which are accessible for an adaptive radiation, starting from the colonization of a novel environment. In the case of repeated colonizations of similar environments from the same source population this "flexible stem" hypothesis predicts similar phenotypes to arise in repeated subsequent radiations. The Midas Cichlid (Amphilophus spp.) in Nicaragua has radiated in parallel in several crater-lakes seeded by populations originating from the Nicaraguan Great Lakes. Here, we tested phenotypic plasticity in the pharyngeal jaw of Midas Cichlids. The pharyngeal jaw apparatus of cichlids, a second set of jaws functionally decoupled from the oral ones, is known to mediate ecological specialization and often differs strongly between sister-species. Results We performed a common garden experiment raising three groups of Midas cichlids on food differing in hardness and calcium content. Analyzing the lower pharyngeal jaw-bones we find significant differences between diet groups qualitatively resembling the differences found between specialized species. Observed differences in pharyngeal jaw expression between groups were attributable to the diet's mechanical resistance, whereas surplus calcium in the diet was not found to be of importance. Conclusions The pharyngeal jaw apparatus of Midas Cichlids can be expressed plastically if stimulated mechanically during feeding. Since this trait is commonly differentiated - among other traits - between
Boileau, Nicolas; Cortesi, Fabio; Egger, Bernd; Muschick, Moritz; Indermaur, Adrian; Theis, Anya; Büscher, Heinz H; Salzburger, Walter
Aggressive mimicry is an adaptive tactic of parasitic or predatory species that closely resemble inoffensive models in order to increase fitness via predatory gains. Although similarity of distantly related species is often intuitively implicated with mimicry, the exact mechanisms and evolutionary causes remain elusive in many cases. Here, we report a complex aggressive mimicry strategy in Plecodus straeleni, a scale-eating cichlid fish from Lake Tanganyika, which imitates two other cichlid species. Employing targeted sequencing on ingested scales, we show that P. straeleni does not preferentially parasitize its models but—contrary to prevailing assumptions—targets a variety of co-occurring dissimilar looking fish species. Combined with tests for visual resemblance and visual modelling from a prey perspective, our results suggest that complex interactions among different cichlid species are involved in this mimicry system.
Boileau, Nicolas; Cortesi, Fabio; Egger, Bernd; Muschick, Moritz; Indermaur, Adrian; Theis, Anya; Büscher, Heinz H.; Salzburger, Walter
Aggressive mimicry is an adaptive tactic of parasitic or predatory species that closely resemble inoffensive models in order to increase fitness via predatory gains. Although similarity of distantly related species is often intuitively implicated with mimicry, the exact mechanisms and evolutionary causes remain elusive in many cases. Here, we report a complex aggressive mimicry strategy in Plecodus straeleni, a scale-eating cichlid fish from Lake Tanganyika, which imitates two other cichlid species. Employing targeted sequencing on ingested scales, we show that P. straeleni does not preferentially parasitize its models but—contrary to prevailing assumptions—targets a variety of co-occurring dissimilar looking fish species. Combined with tests for visual resemblance and visual modelling from a prey perspective, our results suggest that complex interactions among different cichlid species are involved in this mimicry system. PMID:26399975
Adeogun, Aina O; Ibor, Oju R; Adeduntan, Sherifat D; Arukwe, Augustine
The objectives of this study were to develop and validate biomarker techniques for aquatic environmental monitoring of endocrine disrupting chemicals (EDCs) in Nigeria aquatic ecosystems, using the Eleyele Lake, which is a major source of domestic water supply to Ibadan and its surrounding towns, as a model aquatic environment and Tilapia guineensis, as a model organism. A total of 55 male and 28 female fish were used for this study. No significant difference in condition factor was observed between the sexes. Evaluation of gross gonadal morphology of the sampled fish showed 33% intersex prevalence in the sampled population, of which respective 71 and 29% were males and females, with visible testis and ovary developing alongside phenotypic females and males. Plasma concentrations of luteinizing hormone (LH), follicle-stimulating hormone (FSH), 11-ketotestosterone (11-KT) and 17β-estradiol (E2) were performed, showing that male fish had significantly higher plasma LH and E2 concentrations, compared to females. Vitellogenin (Vtg) and zona radiata proteins (Zrp) mRNA levels were significantly higher in males, compared to female fish. Contaminant analysis revealed that PCB 81, 123, 138 and 196 were the only PCB congeners detected in sediment and fish muscle (PCB153 in sediment), while dieldrin was the only organochlorine compound (OC) detected in Eleyele sediment. These responses were used in a multivariate analysis, showing that two principal components were extracted and accounted for 74% of total variation in the dataset. The principal component analysis (PCA) showed that male fish variables were positively correlated with PCB congeners 18 and 123, while female fish showed positive correlations with congener 81, 138, 189, 196, indicating sex-specific pattern of association between PCBs concentrations and biomarker expression. In addition, strong positive correlation between male fish and LH, E2, FSH and Vtg was observed, while female fish positively correlated with
Katangwe, Thembi; Purcell, Janet; Bar-Zeev, Naor; Denis, Brigitte; Montgomery, Jacqui; Alaerts, Maaike; Heyderman, Robert Simon; Dance, David A.B.; Kennedy, Neil; Feasey, Nicholas
A case of human melioidosis caused by a novel sequence type of Burkholderia pseudomallei occurred in a child in Malawi, southern Africa. A literature review showed that human cases reported from the continent have been increasing. PMID:23735189
Salzburger, Walter; Meyer, Axel
With more than 3,000 species, the fish family Cichlidae is one of the most species-rich families of vertebrates. Cichlids occur in southern and central America, Africa, Madagascar, and India. The hotspot of their biodiversity is East Africa, where they form adaptive radiations composed of hundreds of endemic species in several lakes of various sizes and ages. The unparalleled species richness of East African cichlids has been something of a conundrum for evolutionary biologists and ecologists, since it has been in doubt whether these hundreds of species arose by allopatric speciation or whether it is necessary to invoke somewhat less traditional models of speciation, such as micro-allopatric, peripatric, or even sympatric speciation or evolution through sexual selection mediated by female choice. Ernst Mayr's analyses of these evolutionary uniquely diverse species assemblages have contributed to a more direct approach to this problem and have led to a deeper understanding of the patterns and processes that caused the formation of these huge groups of species. We review here recent molecular data on population differentiation and phylogenetics, which have helped to unravel, to some extent, the patterns and processes that led to the formation and ecological maintenance of cichlid species flocks. It is becoming apparent that sexually selected traits do play an important role in speciation in micro-allopatric or even sympatric settings. Species richness seems to be roughly correlated with the surface area, but not the age, of the lakes. We observe that the oldest lineages of a species flock of cichlids are often less species-rich and live in the open water or deepwater habitats. While the species flocks of the Lake Malawai and the Lake Victoria areas were shown to be monophyletic, the cichlid assemblage of Lake Tanganyika seems to consist of several independent species flocks. Cichlids emerge as an evolutionary model system in which many fundamental questions in
Gennaro, Eugene D.; Winters, Charlotte M.
A classroom experiment is proposed in which students can mate a banded or spotted convict cichlid with a pink convict cichlid and observe the markings of their "children" and "grandchildren" as a way of illustrating Mendel's Laws of Dominance and Segregation. (MN)
Malawi lies in Africa's Great Rift Valley. Its western border is defined by Lake Malawi, the third largest lake in Africa. Over 80% of Malawians live in rural areas and 90% of the labor force is associated with agriculture. More than half of the population lives below the poverty line. Area characteristics indicate a high likelihood of nitrate and total coliform in community drinking water. Infants exposed to high nitrate are at risk of developing methemoglobinemia. In addition, diarrheal diseases from unsafe drinking water are one of the top causes of mortality in children under five. Without sufficient and sustainable supplies of clean water, these challenges will continue to threaten Malawi's ability to overcome the devastating impact of diarrheal diseases on its population. Therefore, Malawi remains highly dependent on outside assistance and influence to reduce or eliminate the threat posed by unsafe drinking water. This research presents a literature review of nitrate and total coliform groundwater quality and a proposed risk communication plan for drinking water in northern Malawi.
Stelkens, Rike Bahati; Schmid, Corinne; Seehausen, Ole
Studies from a wide diversity of taxa have shown a negative relationship between genetic compatibility and the divergence time of hybridizing genomes. Theory predicts the main breakdown of fitness to happen after the F1 hybrid generation, when heterosis subsides and recessive allelic (Dobzhansky-Muller) incompatibilities are increasingly unmasked. We measured the fitness of F2 hybrids of African haplochromine cichlid fish bred from species pairs spanning several thousand to several million years divergence time. F2 hybrids consistently showed the lowest viability compared to F1 hybrids and non-hybrid crosses (crosses within the grandparental species), in agreement with hybrid breakdown. Especially the short- and long-term survival (2 weeks to 6 months) of F2 hybrids was significantly reduced. Overall, F2 hybrids showed a fitness reduction of 21% compared to F1 hybrids, and a reduction of 43% compared to the grandparental, non-hybrid crosses. We further observed a decrease of F2 hybrid viability with the genetic distance between grandparental lineages, suggesting an important role for negative epistatic interactions in cichlid fish postzygotic isolation. The estimated time window for successful production of F2 hybrids resulting from our data is consistent with the estimated divergence time between the multiple ancestral lineages that presumably hybridized in three major adaptive radiations of African cichlids. PMID:25996870
Mulder, Monique Borgerhoff; Msalu, Lameck; Caro, Tim; Salerno, Jonathan
Livingstone's second mission site on the shore of Lake Malawi suffers very high rates of consequential lightning strikes. Comprehensive interviewing of victims and their relatives in seven Traditional Authorities in Nkhata Bay District, Malawi revealed that the annual rate of consequential strikes was 419/million, more than six times higher than that in other developing countries; the rate of deaths from lightning was 84/million/year, 5.4 times greater than the highest ever recorded. These remarkable figures reveal that lightning constitutes a significant stochastic source of mortality with potential life history consequences, but it should not deflect attention away from the more prominent causes of mortality in this rural area.
Meier, Joana I.; Marques, David A.; Mwaiko, Salome; Wagner, Catherine E.; Excoffier, Laurent; Seehausen, Ole
Understanding why some evolutionary lineages generate exceptionally high species diversity is an important goal in evolutionary biology. Haplochromine cichlid fishes of Africa's Lake Victoria region encompass >700 diverse species that all evolved in the last 150,000 years. How this ‘Lake Victoria Region Superflock' could evolve on such rapid timescales is an enduring question. Here, we demonstrate that hybridization between two divergent lineages facilitated this process by providing genetic variation that subsequently became recombined and sorted into many new species. Notably, the hybridization event generated exceptional allelic variation at an opsin gene known to be involved in adaptation and speciation. More generally, differentiation between new species is accentuated around variants that were fixed differences between the parental lineages, and that now appear in many new combinations in the radiation species. We conclude that hybridization between divergent lineages, when coincident with ecological opportunity, may facilitate rapid and extensive adaptive radiation. PMID:28186104
Ngwira, Bagrey MM; Tambala, Phillimon; Perez, A Maria; Bowie, Cameron; Molyneux, David H
Mapping distribution of lymphatic filariasis (LF) is a prerequisite for planning national elimination programmes. Results from a nation wide mapping survey for lymphatic filariasis (LF) in Malawi are presented. Thirty-five villages were sampled from 23 districts excluding three districts (Karonga, Chikwawa and Nsanje) that had already been mapped and Likoma, an Island, where access was not possible in the time frame of the survey. Antigenaemia prevalence [based on immunochromatographic card tests (ICT)] ranged from 0% to 35.9%. Villages from the western side of the country and distant from the lake tended to be of lower prevalence. The exception was a village in Mchinji district on the Malawi-Zambia border where a prevalence of 18.2% was found. In contrast villages from lake shore districts [Salima, Mangochi, Balaka and Ntcheu (Bwanje valley)] and Phalombe had prevalences of over 20%. A national map is developed which incorporates data from surveys in Karonga, Chikwawa and Nsanje districts, carried out in 2000. There is a marked decline in prevalence with increasing altitude. Further analysis revealed a strong negative correlation (R2 = 0.7 p < 0.001) between altitude and prevalence. These results suggest that the lake shore, Phalombe plain and the lower Shire valley will be priority areas for the Malawi LF elimination programme. Implications of these findings as regards implementing a national LF elimination programme in Malawi are discussed. PMID:18047646
Jørstad, Hanne; Webersik, Christian
In recent years, research on climate change and human security has received much attention among policy makers and academia alike. Communities in the Global South that rely on an intact resource base and struggle with poverty, existing inequalities and historical injustices will especially be affected by predicted changes in temperature and precipitation. The objective of this article is to better understand under what conditions local communities can adapt to anticipated impacts of climate change. The empirical part of the paper answers the question as to what extent local women engaged in fish processing in the Chilwa Basin in Malawi have experienced climate change and how they are affected by it. The article assesses an adaptation project designed to make those women more resilient to a warmer and more variable climate. The research results show that marketing and improving fish processing as strategies to adapt to climate change have their limitations. The study concludes that livelihood diversification can be a more effective strategy for Malawian women to adapt to a more variable and unpredictable climate rather than exclusively relying on a resource base that is threatened by climate change.
Amcoff, M; Gonzalez-Voyer, A; Kolm, N
Sexual selection has been suggested to be an important driver of speciation in cichlid fishes of the Great Lakes of Africa, and the presence of male egg dummies is proposed to have played a key role. Here, we investigate how mouthbrooding and egg dummies have evolved in Tanganyikan cichlids, the lineage which seeded the other African radiations, with a special emphasis on the egg dummies. Using modern phylogenetic comparative analyses and a phylogeny including 86% of the 200 described species, we provide formal evidence demonstrating correlated evolution between mouthbrooding and egg dummies in Tanganyikan cichlids. These results concur with existing evidence, suggesting that egg dummies have evolved through sensory exploitation. We also demonstrate that there is a strong evolutionary correlation between the presence of egg dummies and both pre- and post-copulatory sexual selection. Moreover, egg dummy evolution was contingent on the intensity of pre- and post-copulatory sexual selection in Tanganyikan cichlids. In sum, our results provide evidence supporting the hypothesis of egg dummies evolving through sensory exploitation and highlight the role of sexual selection in favouring the evolution and maintenance of this trait.
Schwarzer, Julia; Misof, Bernhard; Ifuta, Seraphin N; Schliewen, Ulrich K
Most freshwater diversity is arguably located in networks of rivers and streams, but, in contrast to lacustrine systems riverine radiations, are largely understudied. The extensive rapids of the lower Congo River is one of the few river stretches inhabited by a locally endemic cichlid species flock as well as several species pairs, for which we provide evidence that they have radiated in situ. We use more that 2,000 AFLP markers as well as multilocus sequence datasets to reconstruct their origin, phylogenetic history, as well as the timing of colonization and speciation of two Lower Congo cichlid genera, Steatocranus and Nanochromis. Based on a representative taxon sampling and well resolved phylogenetic hypotheses we demonstrate that a high level of riverine diversity originated in the lower Congo within about 5 mya, which is concordant with age estimates for the hydrological origin of the modern lower Congo River. A spatial genetic structure is present in all widely distributed lineages corresponding to a trisection of the lower Congo River into major biogeographic areas, each with locally endemic species assemblages. With the present study, we provide a phylogenetic framework for a complex system that may serve as a link between African riverine cichlid diversity and the megadiverse cichlid radiations of the East African lakes. Beyond this we give for the first time a biologically estimated age for the origin of the lower Congo River rapids, one of the most extreme freshwater habitats on earth.
Van Damme, D.; Gautier, A.
In Terminal Pliocene-Early Pleistocene times, part of the Malawi Basin was occupied by palaeo-lake Chiwondo. Molluscan biostratigraphy situates this freshwater lake either in the East African wet phase between 2.7-2.4 Ma or that of 2.0-1.8 Ma. In-lake divergent evolution remained restricted to a few molluscan taxa and was very modest. The lacustrine Chiwondo fauna went extinct at the beginning of the Pleistocene. The Modern Lake Malawi malacofauna is poor and descends from ubiquistic South-East African taxa and some Malawi Basin endemics that invaded the present lake after the Late Pleistocene mega-droughts. The Pleistocene aridity crises caused dramatic changes, affecting the malacofauna of all East African lakes. All lacustrine endemic faunas that had evolved in the Pliocene rift lakes, such as palaeo-lake Chiwondo, became extinct. In Lake Tanganyika, the freshwater ecosystem did not crash as in other lakes, but the environmental changes were sufficiently important to trigger a vast radiation. All African endemic lacustrine molluscan clades that are the result of in-lake divergence are hence geologically young, including the vast Lavigeria clade in Lake Tanganyika (ca. 43 species).
Fernald, Russell D.
Cichlid fishes represent one of the most species-rich and rapid radiations of a vertebrate family. These ∼2200 species, predominantly found in the East African Great Lakes, exhibit dramatic differences in anatomy, physiology, and behavior. However, the genetic bases for this radiation, and for the control of their divergent traits, are unknown. A flood of genomic and transcriptomic data promises to suggest mechanisms underlying the diversity, but transgenic technology will be needed to rigorously test the hypotheses generated. Here we demonstrate the successful use of the Tol2 transposon system to generate transgenic Astatotilapia burtoni, a haplochromine cichlid from Lake Tanganyika, carrying the GFP transgene under the control of the ubiquitous EF1α promoter. The transgene integrates into the genome, is successfully passed through the germline, and the widespread GFP expression pattern is stable across siblings and multiple generations. The stable inheritance and expression patterns indicate that the Tol2 system can be applied to generate A. burtoni transgenic lines. Transgenesis has proven to be a powerful technology for manipulating genes and cells in other model organisms and we anticipate that transgenic A. burtoni and other cichlids will be used to test the mechanisms underlying behavior and speciation. PMID:24204902
Lund, S.; Platzman, E. S.; Johnson, T. C.; Scholz, C. A.; Cohen, A. S.; Russell, J. M.
We are developing a regional pattern of Holocene paleomagnetic secular variation (PSV) from four lakes in East Africa - Lake Turkana (3°N), Lake Victoria (1°S), Lake Tanganyika (5°S), and Lake Malawi (10°S). Detailed paleomagnetic and rock magnetic measurements have been made on two cores from Lake Malawi (9m meters in depth, last ~10,000 years), two cores from Lake Victoria (8 m, last ~8,000 years), 11 cores from Lake Turkana (2-9 m, last ~10,000 years), and one core from Lake Tanganyika (5 m, last ~5,000 years). Our rock magnetic studies identify significant intervals of magnetic mineral dissolution in Lakes Victoria and Tanganyika making parts of these cores unsuitable for relative paleointensity studies. On the other hand, rock magnetic variability in the Lake Malawi and Lake Turkana cores are stable and correlatable among cores. We have recovered directional secular variation records from Lakes Malawi, Victoria, and Turkana. Millennial-scale inclination and declination features can be correlated among cores at each lake and between lakes. We have also recovered relative paleointensity records from Lakes Malawi and Turkana. More than 20 radiocarbon dates and detailed seismic stratigraphy (Turkana) provide critical added information for correlating and dating the paleomagnetic records.
Lehtonen, Topi K
The coexistence of species with overlapping resource use is often thought to involve only negative fitness effects as a consequence of interspecific competition. Furthermore, the scarce empirical research on positive species interactions has predominantly focused on sessile organisms. Here, I experimentally assessed the effect of close proximity of a potential brood predator and competitor on reproductive success of a neotropical cichlid fish. I demonstrate that convict cichlid (Archocentrus nigrofasciatus) broods have a higher survival rate near territories of the Nicaragua cichlid (Hypsophrys nicaraguensis), and that escape from predation and nest takeovers is the most likely explanation for the decreased offspring mortality.
Van Steenberge, Maarten; Pariselle, Antoine; Huyse, Tine; Volckaert, Filip A. M.; Snoeks, Jos; Vanhove, Maarten P. M.
The unparalleled biodiversity of Lake Tanganyika (Africa) has fascinated biologists for over a century; its unique cichlid communities are a preferred model for evolutionary research. Although species delineation is, in most cases, relatively straightforward, higher-order classifications were shown not to agree with monophyletic groups. Here, traditional morphological methods meet their limitations. A typical example are the tropheine cichlids currently belonging to Simochromis and Pseudosimochromis. The affiliations of these widespread and abundant cichlids are poorly understood. Molecular work suggested that genus and species boundaries should be revised. Moreover, previous morphological results indicated that intraspecific variation should be considered to delineate species in Lake Tanganyika cichlids. We review the genera Simochromis and Pseudosimochromis using an integrative approach. Besides a morphometric study and a barcoding approach, monogenean Cichlidogyrus (Platyhelminthes: Ancyrocephalidae) gill parasites, often highly species-specific, are used as complementary markers. Six new species are described. Cichlidogyrus raeymaekersi sp. nov., C. muterezii sp. nov. and C. banyankimbonai sp. nov. infect S. diagramma. Cichlidogyrus georgesmertensi sp. nov. was found on S. babaulti and S. pleurospilus, C. franswittei sp. nov. on both S. marginatus and P. curvifrons and C. frankwillemsi sp. nov. only on P. curvifrons. As relatedness between Cichlidogyrus species usually reflects relatedness between hosts, we considered Simochromis monotypic because the three Cichlidogyrus species found on S. diagramma belonged to a different morphotype than those found on the other Simochromis. The transfer of S. babaulti, S. marginatus, S. pleurospilus and S. margaretae to Pseudosimochromis was justified by the similarity of their Cichlidogyrus fauna and the intermediate morphology of S. margaretae. Finally parasite data also supported the synonymy between S. pleurospilus and S
Background Cichlid fishes have been the subject of increasing scientific interest because of their rapid adaptive radiation which has led to an extensive ecological diversity and their enormous importance to tropical and subtropical aquaculture. To increase our understanding of chromosome evolution among cichlid species, karyotypes of one Asian, 22 African, and 30 South American cichlid species were investigated, and chromosomal data of the family was reviewed. Results Although there is extensive variation in the karyotypes of cichlid fishes (from 2n = 32 to 2n = 60 chromosomes), the modal chromosome number for South American species was 2n = 48 and the modal number for the African ones was 2n = 44. The only Asian species analyzed, Etroplus maculatus, was observed to have 46 chromosomes. The presence of one or two macro B chromosomes was detected in two African species. The cytogenetic mapping of 18S ribosomal RNA (18S rRNA) gene revealed a variable number of clusters among species varying from two to six. Conclusions The karyotype diversification of cichlids seems to have occurred through several chromosomal rearrangements involving fissions, fusions and inversions. It was possible to identify karyotype markers for the subfamilies Pseudocrenilabrinae (African) and Cichlinae (American). The karyotype analyses did not clarify the phylogenetic relationship among the Cichlinae tribes. On the other hand, the two major groups of Pseudocrenilabrinae (tilapiine and haplochromine) were clearly discriminated based on the characteristics of their karyotypes. The cytogenetic mapping of 18S ribosomal RNA (18S rRNA) gene did not follow the chromosome diversification in the family. The dynamic evolution of the repeated units of rRNA genes generates patterns of chromosomal distribution that do not help follows the phylogenetic relationships among taxa. The presence of B chromosomes in cichlids is of particular interest because they may not be represented in the reference genome
Singogo, P P; Chintengo, S
Traditional health practitioners (THP) far outnumber biomedical staff in Malawi. They are generally well-known, accepted, and trusted by villagers, and are respected guardians of sociocultural values and opinion leaders concerning health. Many treat 5-6 sexually transmitted disease patients per day. The National AIDS Program therefore enlisted the collaboration of the Herbalist Association of Malawi in a special AIDS project. Objectives were to train THPs as peer educators, increase biomedical practitioners' awareness of traditional healing practices, and promote greater communication and cooperation between THPs and the biomedical sector. Although many THPs are illiterate, their generally literate secretaries help facilitate use of the manual and the dissemination of information. The first training course was organized for THPs from two districts in 1992 with seven more following to train a total 234 THPs. Biomedical practitioners from the district provide support. A participatory approach is used to engage THPs in exercises and group discussions. Follow-up has shown success with THPs understanding that AIDS is incurable and that is has a big impact upon the community. 91% were conducting AIDS education sessions, several have trained other healers in their districts, and some actively distribute condoms in their communities. The project now wants to train traditional birth attendants and involve biomedical health staff in record keeping for the THPs on condom distribution.
Sefc, Kristina M.; Brown, Alexandria C.; Clotfelter, Ethan D.
Animal colors play important roles in communication, ecological interactions and speciation. Carotenoid pigments are responsible for many yellow, orange and red hues in animals. Whereas extensive knowledge on the proximate mechanisms underlying carotenoid coloration in birds has led to testable hypotheses on avian color evolution and signaling, much less is known about the expression of carotenoid coloration in fishes. Here, we promote cichlid fishes (Perciformes: Cichlidae) as a system in which to study the physiological and evolutionary significance of carotenoids. Cichlids include some of the best examples of adaptive radiation and color pattern diversification in vertebrates. In this paper, we examine fitness correlates of carotenoid pigmentation in cichlids and review hypotheses regarding the signal content of carotenoid-based ornaments. Carotenoid-based coloration is influenced by diet and body condition and is positively related to mating success and social dominance. Gaps in our knowledge are discussed in the last part of this review, particularly in the understanding of carotenoid metabolism pathways and the genetics of carotenoid coloration. We suggest that carotenoid metabolism and transport are important proximate mechanisms responsible for individual and population-differences in cichlid coloration that may ultimately contribute to diversification and speciation. PMID:24667558
Hulsey, C Darrin
Cichlid fishes provide an excellent natural system for integrating studies of genomics and adaptive radiation. Cichlids are unique in comprising a substantial fraction of all vertebrate species, possessing unique jaw structures, displaying an exceptional range of breeding systems, and exhibiting rampant phenotypic convergence. The rate of divergence in cichlid jaws, teeth, color patterns, visual systems, reproductive biology, and mating behaviors is unparalleled among vertebrates. I discuss ways rapid divergence in cichlids and other adaptive radiations make understanding the genomic basis of adaptive divergence more tractable. Then, I briefly overview some major findings and insights into vertebrate adaptation that have been gained through cichlid genetic studies. Finally, I discuss the extensive evolutionary replication provided by cichlid adaptive radiations and their potential for studies of genotype-to-phenotype mapping.
Baldo, Laura; Riera, Joan Lluís; Tooming-Klunderud, Ave; Albà, M. Mar; Salzburger, Walter
The gut microbiota structure reflects both a host phylogenetic history and a signature of adaptation to the host ecological, mainly trophic niches. African cichlid fishes, with their array of closely related species that underwent a rapid dietary niche radiation, offer a particularly interesting system to explore the relative contribution of these two factors in nature. Here we surveyed the host intra- and interspecific natural variation of the gut microbiota of five cichlid species from the monophyletic tribe Perissodini of lake Tanganyika, whose members transitioned from being zooplanktivorous to feeding primarily on fish scales. The outgroup riverine species Astatotilapia burtoni, largely omnivorous, was also included in the study. Fusobacteria, Firmicutes and Proteobacteria represented the dominant components in the gut microbiota of all 30 specimens analysed according to two distinct 16S rRNA markers. All members of the Perissodini tribe showed a homogenous pattern of microbial alpha and beta diversities, with no significant qualitative differences, despite changes in diet. The recent diet shift between zooplantkon- and scale-eaters simply reflects on a significant enrichment of Clostridium taxa in scale-eaters where they might be involved in the scale metabolism. Comparison with the omnivorous species A. burtoni suggests that, with increased host phylogenetic distance and/or increasing herbivory, the gut microbiota begins differentiating also at qualitative level. The cichlids show presence of a large conserved core of taxa and a small set of core OTUs (average 13–15%), remarkably stable also in captivity, and putatively favoured by both restricted microbial transmission among related hosts (putatively enhanced by mouthbrooding behavior) and common host constraints. This study sets the basis for a future large-scale investigation of the gut microbiota of cichlids and its adaptation in the process of the host adaptive radiation. PMID:25978452
Fan, Shaohua; Meyer, Axel
African cichlid fishes are an ideal system for studying explosive rates of speciation and the origin of diversity in adaptive radiation. Within the last few million years, more than 2000 species have evolved in the Great Lakes of East Africa, the largest adaptive radiation in vertebrates. These young species show spectacular diversity in their coloration, morphology and behavior. However, little is known about the genomic basis of this astonishing diversity. Recently, five African cichlid genomes were sequenced, including that of the Nile Tilapia (Oreochromis niloticus), a basal and only relatively moderately diversified lineage, and the genomes of four representative endemic species of the adaptive radiations, Neolamprologus brichardi, Astatotilapia burtoni, Metriaclima zebra, and Pundamila nyererei. Using the Tilapia genome as a reference genome, we generated a high-resolution genomic variation map, consisting of single nucleotide polymorphisms (SNPs), short insertions and deletions (indels), inversions and deletions. In total, around 18.8, 17.7, 17.0, and 17.0 million SNPs, 2.3, 2.2, 1.4, and 1.9 million indels, 262, 306, 162, and 154 inversions, and 3509, 2705, 2710, and 2634 deletions were inferred to have evolved in N. brichardi, A. burtoni, P. nyererei, and M. zebra, respectively. Many of these variations affected the annotated gene regions in the genome. Different patterns of genetic variation were detected during the adaptive radiation of African cichlid fishes. For SNPs, the highest rate of evolution was detected in the common ancestor of N. brichardi, A. burtoni, P. nyererei, and M. zebra. However, for the evolution of inversions and deletions, we found that the rates at the terminal taxa are substantially higher than the rates at the ancestral lineages. The high-resolution map provides an ideal opportunity to understand the genomic bases of the adaptive radiation of African cichlid fishes. PMID:24917883
Frickey, Tancred; Jones, Julia C.; Meyer, Axel
Gut bacterial communities are now known to influence a range of fitness related aspects of organisms. But how different the microbial community is in closely related species, and if these differences can be interpreted as adaptive is still unclear. In this study we compared microbial communities in two sets of closely related sympatric crater lake cichlid fish species pairs that show similar adaptations along the limnetic-benthic axis. The gut microbial community composition differs in the species pair inhabiting the older of two crater lakes. One major difference, relative to other fish, is that in these cichlids that live in hypersaline crater lakes, the microbial community is largely made up of Oceanospirillales (52.28%) which are halotolerant or halophilic bacteria. This analysis opens up further avenues to identify candidate symbiotic or co-evolved bacteria playing a role in adaptation to similar diets and life-styles or even have a role in speciation. Future functional and phylosymbiotic analyses might help to address these issues. PMID:24733403
Wien, Carol Anne
The lake is blue black and deep. It is a glaciated finger lake, clawed out of rock when ice retracted across Nova Scotia in a northerly direction during the last ice age. The lake is narrow, a little over a mile long, and deep, 90 to 190 feet in places according to local lore, off the charts in others. The author loves to swim there, with a sense…
Nkhoma, Bryson G.; Mulwafu, Wapulumuka O.
This study examines the process of irrigation management transfer (IMT) in Malawi using the case studies of Likangala and Domasi irrigation schemes in the lake Chilwa basin. It observes that the concept of irrigation management transfer has been adopted against the background of poor performance of irrigation schemes in Malawi and the desire of the state to conform to global trends and approaches in the management of irrigation schemes. In the context of Malawi, the process of IMT involves rehabilitation of schemes and training of farmers before handing them over. The paper notes that so far the handover process has been fraught with many challenges, not least of which includes lack of adequate funds, delays, lack of public awareness and farmers’ conflicting perceptions on the handover of irrigation schemes. These problems militate against the success of IMT in the Domasi and Likangala irrigation schemes in particular and other schemes in the country as a whole.
The Drought Network for information sharing eventually led to the establishment of the more formal Southern Region AIDS Network (SORAN) where representatives from government and nongovernmental organizations (NGOs) focused on awareness raising, lobbying, and advocacy. As an initial step towards networking on HIV/AIDS issues, a festival was organized in Blantyre on December 4, 1993, by NGOs, private companies, church groups, school children, and volunteers to bring about behavior change. About 2000 people gathered to listen to music, learn about HIV transmission through drama group presentations, watch videos with HIV/STD prevention messages, and learn about proper condom use. The participants officially established SORAN in February 1994 to act as a coordinating body for organizations working in prevention and care for HIV/STD-infected persons and their families. Network activities endeavored: to assist organizations interested in developing HIV/AIDS programs and activities; to encourage the business communities to participate in multisectoral coordination and to help channel funds from them to HIV/AIDS programs; to act as a resource center for information about HIV/AIDS; and to lobby among politicians as well as traditional local and religious leaders. When the first multi-party parliamentary election approached in May 1994, SORAN challenged representatives of 7 political parties and a women's organization to speak out publicly on what they envisioned doing about HIV/AIDS. The Grand Walk was also organized by SORAN members representing the Catholic Episcopal Conference of Malawi, the Protestant Blantyre Synod, a local brewery, and UNICEF. About 500 walkers received support from passersby. 70% were school children 10-18 years old who sang AIDS awareness songs and passed out flyers. Three months later the National AIDS Program's Big Walk for AIDS, following a National AIDS Crisis Conference, signaled the government's public recognition of the need for a
Lorin, Thibault; Salzburger, Walter; Böhne, Astrid
The emergence of the steroid system is coupled to the evolution of multicellular animals. In vertebrates in particular, the steroid receptor repertoire has been shaped by genome duplications characteristic to this lineage. Here, we investigate for the first time the composition of the androgen receptor–signaling pathway in ray-finned fish genomes by focusing in particular on duplicates that emerged from the teleost-specific whole-genome duplication. We trace lineage- and species-specific duplications and gene losses for the genomic and nongenomic pathway of androgen signaling and subsequently investigate the sequence evolution of these genes. In one particular fish lineage, the cichlids, we find evidence for differing selection pressures acting on teleost-specific whole-genome duplication paralogs at a derived evolutionary stage. We then look into the expression of these duplicated genes in four cichlid species from Lake Tanganyika indicating, once more, rapid changes in expression patterns in closely related fish species. We focus on a particular case, the cichlid specific duplication of the rac1 GTPase, which shows possible signs of a neofunctionalization event. PMID:26333839
Nikaido, Masato; Suzuki, Hikoyu; Toyoda, Atsushi; Fujiyama, Asao; Hagino-Yamagishi, Kimiko; Kocher, Thomas D.; Carleton, Karen; Okada, Norihiro
Fish use olfaction to sense a variety of nonvolatile chemical signals in water. However, the evolutionary importance of olfaction in species-rich cichlids is controversial. Here, we determined an almost complete sequence of the vomeronasal type 2 receptor-like (OlfC: putative amino acids receptor in teleosts) gene cluster using the bacterial artificial chromosome library of the Lake Victoria cichlid, Haplochromis chilotes. In the cluster region, we found 61 intact OlfC genes, which is the largest number of OlfC genes identified among the seven teleost fish investigated to date. Data mining of the Oreochromis niloticus (Nile tilapia) draft genome sequence, and genomic Southern hybridization analysis revealed that the ancestor of all modern cichlids had already developed almost the same OlfC gene repertoire, which was accomplished by lineage-specific gene expansions. Furthermore, comparison of receptor sequences showed that recently duplicated paralogs are more variable than orthologs of different species at particular sites that were predicted to be involved in amino acid selectivity. Thus, the increase of paralogs through gene expansion may lead to functional diversification in detection of amino acids. This study implies that cichlids have developed a potent capacity to detect a variety of amino acids (and their derivatives) through OlfCs, which may have contributed to the extraordinary diversity of their feeding habitats. PMID:23501830
Hulsey, C Darrin; García-de-León, Francisco J
Trophically polymorphic species could represent lineages that are rapidly diverging along an ecological axis or could phenotypically mark the collapse of species through introgressive hybridization. We investigated patterns of introgression between the trophically polymorphic cichlid fish Herichthys minckleyi and its relative H. cyanoguttatus using a combination of population genetics and species tree analyses. We first examined the distribution of mitochondrial haplotypes within the alternative H. minckleyi pharyngeal jaw morphotypes that are endemic to the small desert valley of Cuatro Ciénegas. We recovered two clusters of mitochondrial haplotypes. The first contained a number of slightly differentiated cytochrome b (cytb) haplotypes that showed some phylogeographic signal and were present in both jaw morphotypes. The other haplotype was monomorphic, highly differentiated from the other cluster, present in equal frequencies in the morphotypes, and identical to H. cyanoguttatus haplotypes found outside Cuatro Ciénegas. Then, we investigated whether H. minckleyi individuals with the H. cyanoguttatus cytb were more evolutionarily similar to H. cyanoguttatus or other H. minckleyi using a species tree analysis of 84 nuclear loci. Both H. minckleyi pharyngeal morphotypes, regardless of their cytb haplotype, were quite distinct from H. cyanoguttatus. However, hybridization could be blurring subdivision within H. minckleyi as the alternative jaw morphotypes were not genetically distinct from one another. Accounting for introgression from H. cyanoguttatus will be essential to understand the evolution of the trophically polymorphic cichlid H. minckleyi. PMID:24340193
Lehtonen, Topi K.; McCrary, Jeffrey K.; Meyer, Axel
Introduced species represent one of the most serious global threats to biodiversity. In this field-based study, we assessed behavioural responses of brood tending cichlid fish to an invasive predator of their offspring. This was achieved by comparing parental defence responses of the endangered arrow cichlid (Amphilophus zaliosus), a fish species endemic to the crater lake Apoyo in Nicaragua, towards the bigmouth sleeper (Gobiomorus dormitor), a formidable predator of cichlid fry, and all other potential fish predators of offspring. The bigmouth sleeper was recently introduced into Apoyo but naturally co-exists with cichlids in a few other Nicaraguan lakes. Arrow cichlid parents allowed bigmouth sleepers to advance much closer to their fry than other predators before initiating aggressive brood defence behaviours. Interestingly, parents of a very closely related species, A. sagittae, which has coevolved with bigmouth sleepers in crater lake Xiloá, reacted to approaching bigmouth sleepers at comparable distances as to other predators of cichlid fry. These results provide a novel demonstration of the specific mechanism (i.e. naive parental behaviour) by which invasive predators may negatively affect species that lack the adequate behavioural repertoire. PMID:22253881
Matemba, Yonah Hisbon
This article gives an overview of (Christian) Theological Education (CTE) in Malawi. To place the discussion in its appropriate context, information about Malawi is given including the impact of Christianity on the country. The article then describes historical aspects of CTE and highlights some of its inherent shortcomings, before shifting to…
Manousaki, Tereza; Hull, Pincelli M; Kusche, Henrik; Machado-Schiaffino, Gonzalo; Franchini, Paolo; Harrod, Chris; Elmer, Kathryn R; Meyer, Axel
The study of parallel evolution facilitates the discovery of common rules of diversification. Here, we examine the repeated evolution of thick lips in Midas cichlid fishes (the Amphilophus citrinellus species complex)-from two Great Lakes and two crater lakes in Nicaragua-to assess whether similar changes in ecology, phenotypic trophic traits and gene expression accompany parallel trait evolution. Using next-generation sequencing technology, we characterize transcriptome-wide differential gene expression in the lips of wild-caught sympatric thick- and thin-lipped cichlids from all four instances of repeated thick-lip evolution. Six genes (apolipoprotein D, myelin-associated glycoprotein precursor, four-and-a-half LIM domain protein 2, calpain-9, GTPase IMAP family member 8-like and one hypothetical protein) are significantly underexpressed in the thick-lipped morph across all four lakes. However, other aspects of lips' gene expression in sympatric morphs differ in a lake-specific pattern, including the magnitude of differentially expressed genes (97-510). Generally, fewer genes are differentially expressed among morphs in the younger crater lakes than in those from the older Great Lakes. Body shape, lower pharyngeal jaw size and shape, and stable isotopes (δ(13)C and δ(15)N) differ between all sympatric morphs, with the greatest differentiation in the Great Lake Nicaragua. Some ecological traits evolve in parallel (those related to foraging ecology; e.g. lip size, body and head shape) but others, somewhat surprisingly, do not (those related to diet and food processing; e.g. jaw size and shape, stable isotopes). Taken together, this case of parallelism among thick- and thin-lipped cichlids shows a mosaic pattern of parallel and nonparallel evolution.
Balshine, Sigal; Wong, Marian Y L; Reddon, Adam R
Even closely related and ecologically similar cichlid species of Lake Tanganyika exhibit an impressive diversity of social systems, and therefore these fishes offer an excellent opportunity to examine the evolution of social behaviour. Sophisticated social relationships are thought to have evolved via a building block design where more fundamental social behaviours and cognitive processes have been combined, incrementally modified, and elaborated over time. Here, we studied two of these putative social building blocks in two closely related species of cichlids: Neolamprologus pulcher, a group-living species, and Telmatochromis temporalis, a non-grouping species. Otherwise well matched in ecology, this pair of species provide an excellent comparison point to understand how behavioural processes may have been modified in relation to the evolution of sociality. Using social assays in both the laboratory and in the field, we explored each species' motivation to interact with conspecifics, and each species' conflict resolution tactics. We found that individuals of the group living species, N. pulcher, displayed higher social motivation and were more likely to produce submission displays than were individuals of the non-grouping species, T. temporalis. We argue that the motivation to interact with conspecifics is a necessary prerequisite for the emergence of group living, and that the use of submission reduces the costs of conflict and facilitates the maintenance of close social proximity. These results suggest that social motivation and conflict resolution tactics are associated with social complexity, and that these behavioural traits may be functionally significant in the evolution and maintenance of sociality.
Schaedelin, Franziska C; van Dongen, Wouter F D; Wagner, Richard H
Parental care of unrelated offspring is widespread but not well understood. We used 11 polymorphic microsatellite loci to investigate the relatedness of fry and parentally caring adults in a 118-nest colony of the socially and genetically monogamous cichlid fish Neolamprologus caudopunctatus in Lake Tanganyika. There was a high proportion of brood mixing, with 59% of 32 broods containing fry unrelated to both parents, and 18% of all 291 sampled fry being unrelated to the breeding pair. There was no evidence of kin selection for adoption because the genetic and foster parents were not more related than expected by chance. Parentage was assigned to 12 adopted fry from 10 broods. Distances traversed by fry varied markedly, from less than one to over 40 meters. The larger distances suggest that at least some brood mixing was instigated by parents transporting portions of their broods in their mouths, as occurs in some cichlids. Further evidence of non-random brood mixing was that foreign fry did not differ in size from their foster siblings within broods, even though they were significantly larger than fry produced by the tending pairs within the colony. These findings suggest that at least some foreign fry had dispersed non-randomly and were adopted by their foster parents. Enlarged broods are known to provide reduced per capita predation, making it potentially adaptive for breeders to adopt unrelated offspring.
Nevado, B; Mautner, S; Sturmbauer, C; Verheyen, E
Understanding how genetic variation is generated and maintained in natural populations, and how this process unfolds in a changing environment, remains a central issue in biological research. In this work, we analysed patterns of genetic diversity from several populations of three cichlid species from Lake Tanganyika in parallel, using the mitochondrial DNA control region. We sampled populations inhabiting the littoral rocky habitats in both very deep and very shallow areas of the lake. We hypothesized that the former would constitute relatively older, more stable and genetically more diverse populations, because they should have been less severely affected by the well-documented episodes of dramatic water-level fluctuations. In agreement with our predictions, populations of all three species sampled in very shallow shorelines showed traces of stronger population growth than populations of the same species inhabiting deep shorelines. However, contrary to our working hypothesis, we found a significant trend towards increased genetic diversity in the younger, demographically less stable populations inhabiting shallow areas, in comparison with the older and more stable populations inhabiting the deep shorelines. We interpret this finding as the result of the establishment of metapopulation dynamics in the former shorelines, by the frequent perturbation and reshuffling of individuals between populations due to the lake-level fluctuations. The repeated succession of periods of allopatric separation and secondary contact is likely to have further increased the rapid pace of speciation in lacustrine cichlids. PMID:23837841
Tsuboi, Masahito; Gonzalez-Voyer, Alejandro; Kolm, Niclas
Functional coupling, where a single morphological trait performs multiple functions, is a universal feature of organismal design. Theory suggests that functional coupling may constrain the rate of phenotypic evolution, yet empirical tests of this hypothesis are rare. In fish, the evolutionary transition from guarding the eggs on a sandy/rocky substrate (i.e. substrate guarding) to mouthbrooding introduces a novel function to the craniofacial system and offers an ideal opportunity to test the functional coupling hypothesis. Using a combination of geometric morphometrics and a recently developed phylogenetic comparative method, we found that head morphology evolution was 43% faster in substrate guarding species than in mouthbrooding species. Furthermore, for species in which females were solely responsible for mouthbrooding the males had a higher rate of head morphology evolution than in those with bi-parental mouthbrooding. Our results support the hypothesis that adaptations resulting in functional coupling constrain phenotypic evolution. PMID:25948565
Bishop, M.G. )
Sedimentary deposits of the Malawi continental rift basin are a direct result of topography and tectonics unique to rift structure. Recent models describe rifts as asymmetric half-graben connected in series by transfer of accommodation zones. Half-graben consist of roughly parallel, tilted fault blocks stepping up from the bounding fault zone where maximum subsidence occurs. The rift becomes a local baselevel and depocenter as regional drainage is shed away by the rift shoulders. Most of the sediments are derived locally due to internal drainage of connected basins, individual basins, and individual fault blocks. The patterns of sedimentation and facies associations depend on structural position at both fault block and half-graben scales. Drainage is directed and dammed by tilted fault blocks. Forward-tilted fault blocks form basinward-thickening sediment wedges filled with facies of axial fluvial systems, alluvial fault-scarp fans, and ponded swamp and lake deposits. These deposits are asymmetrically shifted toward the controlling fault and onlap the upthrown side of the block, ordinarily the site of erosion or nondeposition. Rivers entering the lake on back tilted fault blocks form large deltas resulting in basinward fining and thinning sediment wedges. Lacustrine, nearshore, shoreline, and lake shore plain deposits over multiple fault blocks record lake levels, water chemistry, and tectonic episodes. Tectonic movement periodically changes the basin depth, configuration, and baselevel. This movement results in widespread unconformities deposition and reworking of sediments within the rift.
Roth, R L
An acquired immunodeficiency syndrome (AIDS) prevention project was begun in Malawi in January 1993 by Peace Corps Malawi. 23 workers strive: 1) to offer health education and counseling with regard to human immunodeficiency virus (HIV) infection and AIDS in district hospitals, health centers, and health clinics; 2) to encourage the participation of community groups in AIDS prevention, education, and counseling support activities; and 3) to implement AIDS education in primary and secondary schools. Volunteer activities include health education, home-based care, school programs (drama groups, peer counseling), income-generating activities, condom distribution, and formation of district committees and sub-committees. Target groups include women and youth. Sexual health is promoted. Malawian counterparts are being trained by the volunteers to ensure sustainability; local volunteers are becoming important as resources become scarce. The program is becoming decentralized as it moves into the villages, and community groups are increasing in number. Obstacles include: 1) the lack of resources; 2) the fact that the District AIDS Coordinators are also clinical officers and medical assistants and so can serve only part time; 3) the cultural taboos that make discussion of certain topics difficult; 4) the political system; 5) illiteracy among women; 6) drug abuse among youth; and 7) the difficulty of remaining separated from one's work when one is surrounded by clients, including coworkers, who are HIV positive.
Keck, Benjamin P; Hulsey, C Darrin
The incredibly species-rich cichlid fish faunas of both the Neotropics and Africa are generally thought to be reciprocally monophyletic. However, the phylogenetic affinity of the African cichlid Heterochromis multidens is ambiguous, and this distinct lineage could make African cichlids paraphyletic. In past studies, Heterochromis has been variously suggested to be one of the earliest diverging lineages within either the Neotropical or the African cichlid radiations, and it has even been hypothesized to be the sister lineage to a clade containing all Neotropical and African cichlids. We examined the phylogenetic relationships among a representative sample of cichlids with a dataset of 29 nuclear loci to assess the support for the different hypotheses of the phylogenetic position of Heterochromis. Although individual gene trees in some instances supported alternative relationships, a majority of gene trees, integration of genes into species trees, and hypothesis testing of putative topologies all supported Heterochromis as belonging to the clade of African cichlids.
Lüdecke, Tina; Schrenk, Friedemann; Thiemeyer, Heinrich; Kullmer, Ottmar; Bromage, Timothy G; Sandrock, Oliver; Fiebig, Jens; Mulch, Andreas
The development of East African savannas is crucial for the origin and evolution of early hominins. These ecosystems, however, vary widely in their fraction of woody cover and today range from closed woodland to open grassland savanna. Here, we present the first Plio-Pleistocene long-term carbon isotope (δ(13)C) record from pedogenic carbonate and Suidae teeth in the southern East African Rift (EAR). These δ(13)C data from the Chiwondo and Chitimwe Beds (Karonga Basin, Northern Malawi) represent a southern hemisphere record in the EAR, a key region for reconstructing vegetation patterns in today's Zambezian Savanna, and permit correlation with data on the evolution and migration of early hominins in today's Somali-Masai Endemic Zone. The sediments along the northwestern shore of Lake Malawi contain fossils attributed to Homo rudolfensis and Paranthropus boisei. The associated hominin localities (Uraha, Malema) are situated between the well-known hominin bearing sites of the Somali-Masai Endemic Zone in the Eastern Rift and the Highveld Grassland in southern Africa, and fill an important geographical gap for hominin research. Persistent δ(13)C values around -9‰ from pedogenic carbonate and suid enamel covering the last ∼4.3 Ma indicate a C3-dominated closed environment with regional patches of C4-grasslands in the Karonga Basin. The overall fraction of woody cover of 60-70% reflects significantly higher canopy density in the Malawi Rift than the Eastern Rift through time. The discrepancy between the two savanna types originated in the Late Pliocene, when the Somali-Masai ecosystem started to show increasing evidence for open, C4-dominated landscapes. Based on the Malawi δ(13)C data, the evolution of savanna ecosystems in Eastern Africa followed different patterns along the north-south extent of the EAR. The appearance of C4-grasses is considered a driver of evolutionary faunal shifts, but despite the difference of ecosystem evolution in the north, similar
Bezault, Etienne; Rognon, Xavier; Gharbi, Karim; Baroiller, Jean-Francois; Chevassus, Bernard
The transfer of the genomic resources developed in the Nile tilapia, Oreochromis niloticus, to other Tilapiines sensu lato and African cichlid would provide new possibilities to study this amazing group from genetics, ecology, evolution, aquaculture, and conservation point of view. We tested the cross-species amplification of 32 O. niloticus microsatellite markers in a panel of 15 species from 5 different African cichlid tribes: Oreochromines (Oreochromis, Sarotherodon), Boreotilapiines (Tilapia), Chromidotilapines, Hemichromines, and Haplochromines. Amplification was successfully observed for 29 markers (91%), with a frequency of polymorphic (P(95)) loci per species around 70%. The mean number of alleles per locus and species was 3.2 but varied from 3.7 within Oreochromis species to 1.6 within the nontilapia species. The high level of cross-species amplification and polymorphism of the microsatellite markers tested in this study provides powerful tools for a wide range of molecular genetic studies within tilapia species as well as for other African cichlids.
Ladich, Friedrich; Schulz-Mirbach, Tanja
Background Hearing thresholds of fishes are typically acquired under laboratory conditions. This does not reflect the situation in natural habitats, where ambient noise may mask their hearing sensitivities. In the current study we investigate hearing in terms of sound pressure (SPL) and particle acceleration levels (PAL) of two cichlid species within the naturally occurring range of noise levels. This enabled us to determine whether species with and without hearing specializations are differently affected by noise. Methodology/Principal Findings We investigated auditory sensitivities in the orange chromide Etroplus maculatus, which possesses anterior swim bladder extensions, and the slender lionhead cichlid Steatocranus tinanti, in which the swim bladder is much smaller and lacks extensions. E. maculatus was tested between 0.2 and 3kHz and S. tinanti between 0.1 and 0.5 kHz using the auditory evoked potential (AEP) recording technique. In both species, SPL and PAL audiograms were determined in the presence of quiet laboratory conditions (baseline) and continuous white noise of 110 and 130 dB RMS. Baseline thresholds showed greatest hearing sensitivity around 0.5 kHz (SPL) and 0.2 kHz (PAL) in E. maculatus and 0.2 kHz in S. tinanti. White noise of 110 dB elevated the thresholds by 0–11 dB (SPL) and 7–11 dB (PAL) in E. maculatus and by 1–2 dB (SPL) and by 1–4 dB (PAL) in S. tinanti. White noise of 130 dB elevated hearing thresholds by 13–29 dB (SPL) and 26–32 dB (PAL) in E. maculatus and 6–16 dB (SPL) and 6–19 dB (PAL) in S. tinanti. Conclusions Our data showed for the first time for SPL and PAL thresholds that the specialized species was masked by different noise regimes at almost all frequencies, whereas the non-specialized species was much less affected. This indicates that noise can limit sound detection and acoustic orientation differently within a single fish family. PMID:23469032
thesis are entirely those of the author, and do not represent the official position of the Malawi Defense Force ( MDF ) or Malawi Government...and the long-standing tensions between them, the MYP and MDF erupted into open conflict. On December 3, the MDF attacked MYP headquarters and the... MDF blocked a political rally of UDF. See also Nyasa Times, “Ex-President and Former Malawi Defense Force Commander Urges Malawi Army to Uphold
Schistosomiasis (i.e., "snail fever" or "bilharzia") is a parasitic infection caused by trematodes (flukes) and is endemic in 74 countries in Africa, South America, the Caribbean, and Asia. U.S. residents who work or travel in these countries may be at risk for schistosomiasis. During 1992, two U.S. Peace Corps volunteers (PCVs) were evacuated from Africa because of Schistosoma hematobium infection of the central nervous system (CNS). Both were exposed to fresh water while vacationing at Cape Maclear, a popular resort area on Lake Malawi (Figure 1), in December 1991. This report summarizes the investigation of these two cases and a follow-up investigation of expatriates residing in Malawi.
Málaga-Trillo, E; Zaleska-Rutczynska, Z; McAndrew, B; Vincek, V; Figueroa, F; Sültmann, H; Klein, J
The species flocks of cichlid fishes in the Great East African Lakes are paradigms of adaptive radiation and hence, of great interest to evolutionary biologists. Phylogenetic studies of these fishes have, however, been hampered by the lack of suitable polymorphic markers. The genes of the major histocompatibility complex hold the promise to provide, through their extensive polymorphism, a large number of such markers, but their use has been hampered by the complexity of the genetic system and the lack of definition of the individual loci. In this study we take the first substantial step to alleviate this problem. Using a combination of methods, including the typing of single sperm cells, gyno- or androgenetic individuals, and haploid embryos, as well as sequencing of class II B restriction fragments isolated from gels for Southern blots, we identify the previously characterized homology groups as distinct loci. At least 17 polymorphic class II B loci, all of which are presumably transcribed, have been found among the different species studied. Most of these loci are shared across the various cichlid species and genera. The number of loci per haplotype varies from individual to individual, ranging from 1 to 13. A total of 21 distinct haplotypes differing in the number of loci they carry has thus far been identified. All the polymorphic loci are part of the same cluster in which, however, distances between at least some of the loci (as indicated by recombination frequencies) are relatively large. Both the individual loci and the haplotypes can now be used to study phylogenetic relationships among the members of the species flocks and the mode in which speciation occurs during adaptive radiation. PMID:9649539
Chazema, Jacqueline; O'Meara, James
The establishment of formal teacher training systems is a relatively new innovation in some parts of Africa. Formal training of secondary teachers in Malawi commenced when the government of Malawi opened the University of Malawi in 1965. The contribution of Malawi's ruling parties to the development of teacher education in Malawi since then is an…
Darrin Hulsey, C
The pharyngeal jaw of cichlids may represent a key innovation that facilitated their unparalleled trophic divergence. In cichlids, ‘fusion’ of the lower pharyngeal jaw (LPJ) results from suturing between the two lower ceratobranchials. To examine, what novel abilities a more extensively fused pharyngeal jaw may confer, the function of LPJ suturing was examined in Heroine cichlids. Greater LPJ suturing, pharyngeal jaw splitting under compression and the forces used to crush molluscs in the wild suggest increased LPJ fusion in the trophically polymorphic Herichthys minckleyi operates to strengthen the pharyngeal jaw. Among Heroine cichlid species, the presence of an external LPJ suture and feeding specialization on molluscs was evolutionarily quite variable, but greater LPJ fusion estimated from the amount of external suturing was highly correlated with molluscivory. Throughout cichlid diversification, increased pharyngeal jaw fusion via suturing has likely helped to reinforce the LPJ during pharyngeal processing thereby facilitating the ability of cichlids to exploit durable prey. PMID:16608685
Franchini, Paolo; Monné Parera, Daniel; Kautt, Andreas F; Meyer, Axel
The identification of thousands of variants across the genomes and their accurate genotyping are crucial for estimating the genetic parameters needed to address a host of molecular ecological and evolutionary questions. With rapid advances of massively parallel high-throughput sequencing technologies, several methods have recently been developed to access genomewide data on population variation. One of the most successful and widely used techniques relies on the combination of restriction enzymes and sequencing-by-synthesis: restriction-site-associated DNA sequencing (RADSeq). We developed a new, more time- and cost-efficient double-digest RAD paired-end protocol (quaddRAD) that simplifies and speeds up the identification of PCR duplicates and permits large-scale multiplexing. Assessing its performance on a technical data set, we also applied the quaddRAD method on population samples of a Neotropical cichlid fish lineage (Archocentrus centrarchus) to assess its genetic structure and demographic history. While we identified allopatric interlake genetic divergence, most likely driven by drift, no signature of sympatric divergence was detected. This differs from what has been observed in the clade of Midas cichlids (Amphilophus citrinellus spp.), another cichlid lineage that inhabits the same lakes and shares a similar demographic history, but has evolved into small-scale adaptive radiations via sympatric speciation. We demonstrate that quaddRAD is a robust and efficient method for genotyping a massive number and widely overlapping set of loci with high accuracy. Furthermore, the results on A. centrarchus open new research avenues providing an ideal system to investigate genome-level mechanisms that could alter the speciation potential of different but closely related cichlid lineages.
Colbourn, Tim; Lewycka, Sonia; Nambiar, Bejoy; Anwar, Iqbal; Phoya, Ann; Mhango, Chisale
Background Millennium Development Goal 5 (MDG 5) targets a 75% reduction in maternal mortality from 1990 to 2015, yet accurate information on trends in maternal mortality and what drives them is sparse. We aimed to fill this gap for Malawi, a country in sub-Saharan Africa with high maternal mortality. Methods We reviewed the literature for population-based studies that provide estimates of the maternal mortality ratio (MMR) in Malawi, and for studies that list and justify variables potentially associated with trends in MMR. We used all population-based estimates of MMR representative of the whole of Malawi to construct a best-fit trend-line for the range of years with available data, calculated the proportion attributable to HIV and qualitatively analysed trends and evidence related to other covariates to logically assess likely candidate drivers of the observed trend in MMR. Results 14 suitable estimates of MMR were found, covering the years 1977–2010. The resulting best-fit line predicted MMR in Malawi to have increased from 317 maternal deaths/100 000 live-births in 1980 to 748 in 1990, before peaking at 971 in 1999, and falling to 846 in 2005 and 484 in 2010. Concurrent deteriorations and improvements in HIV and health system investment and provisions are the most plausible explanations for the trend. Female literacy and education, family planning and poverty reduction could play more of a role if thresholds are passed in the coming years. Conclusions The decrease in MMR in Malawi is encouraging as it appears that recent efforts to control HIV and improve the health system are bearing fruit. Sustained efforts to prevent and treat maternal complications are required if Malawi is to attain the MDG 5 target and save the lives of more of its mothers in years to come. PMID:24353257
Valente, Guilherme T; Conte, Matthew A; Fantinatti, Bruno E A; Cabral-de-Mello, Diogo C; Carvalho, Robson F; Vicari, Marcelo R; Kocher, Thomas D; Martins, Cesar
Approximately 15% of eukaryotes contain supernumerary B chromosomes. When present, B chromosomes frequently represent as much as 5% of the genome. Despite thousands of reports describing the distribution of supernumeraries in various taxa, a comprehensive theory for the origin, maintenance, and evolution of B chromosomes has not emerged. Here, we sequence the complete genomes of individual cichlid fish (Astatotilapia latifasciata) with and without B chromosomes, as well as microdissected B chromosomes, to identify DNA sequences on the B. B sequences were further analyzed through quantitative polymerase chain reaction and in situ hybridization. We find that the B chromosome contains thousands of sequences duplicated from essentially every chromosome in the ancestral karyotype. Although most genes on the B chromosome are fragmented, a few are largely intact, and we detect evidence that at least three of them are transcriptionally active. We propose a model in which the B chromosome originated early in the evolutionary history of Lake Victoria cichlids from a small fragment of one autosome. DNA sequences originating from several autosomes, including protein-coding genes and transposable elements, subsequently inserted into this proto-B. We propose that intact B chromosome genes involved with microtubule organization, kinetochore structure, recombination and progression through the cell cycle may play a role in driving the transmission of the B chromosome. Furthermore, our work suggests that karyotyping is an essential step prior to genome sequencing to avoid problems in genome assembly and analytical biases created by the presence of high copy number sequences on the B chromosome.
Mhango, Chisale; Philbin, Jesse; Chimwaza, Wanangwa; Chipeta, Effie; Msusa, Ausbert
Background In Malawi, abortion is legal only if performed to save a woman’s life; other attempts to procure an abortion are punishable by 7–14 years imprisonment. Most induced abortions in Malawi are performed under unsafe conditions, contributing to Malawi’s high maternal mortality ratio. Malawians are currently debating whether to provide additional exceptions under which an abortion may be legally obtained. An estimated 67,300 induced abortions occurred in Malawi in 2009 (equivalent to 23 abortions per 1,000 women aged 15–44), but changes since 2009, including dramatic increases in contraceptive prevalence, may have impacted abortion rates. Methods We conducted a nationally representative survey of health facilities to estimate the number of cases of post-abortion care, as well as a survey of knowledgeable informants to estimate the probability of needing and obtaining post-abortion care following induced abortion. These data were combined with national population and fertility data to determine current estimates of induced abortion and unintended pregnancy in Malawi using the Abortion Incidence Complications Methodology. Results We estimate that approximately 141,044 (95% CI: 121,161–160,928) induced abortions occurred in Malawi in 2015, translating to a national rate of 38 abortions per 1,000 women aged 15–49 (95% CI: 32 to 43); which varied by geographical zone (range: 28–61). We estimate that 53% of pregnancies in Malawi are unintended, and that 30% of unintended pregnancies end in abortion. Given the challenges of estimating induced abortion, and the assumptions required for calculation, results should be viewed as approximate estimates, rather than exact measures. Conclusions The estimated abortion rate in 2015 is higher than in 2009 (potentially due to methodological differences), but similar to recent estimates from nearby countries including Tanzania (36), Uganda (39), and regional estimates in Eastern and Southern Africa (34–35). Over
One of the biggest challenges for healthcare professionals working in developing countries is the lack of trained personnel to carry out much needed health care provision. Shirley Stronge worked as a nurse/midwife tutor in a rural area in the north of Malawi. Millennium Development Goals four and five have focused our attention on the care required by mothers and newborns. Shirley has chosen to reflect on the role of Traditional Birth Attendants in the north of Malawi and their positive impact on maternity services in this area.
In sub-Saharan Africa, health systems are fragile and staffing is grossly inadequate to meet rising health needs. Despite growing international attention, donors have been reluctant to undertake the significant investments required to address the human resources problem comprehensively, given social and political sensitivities, and concerns regarding sustainability of interventions and risks of rising donor dependency. In Malawi, one of the poorest nations in Africa, declining human resource levels have fueled an accelerating collapse of public health services since the late 1990s. In an effort to improve health outcomes, in 2004 the government launched a new health initiative to deliver an Essential Health Package, including a major scale-up of HIV and AIDS related services. Improving staffing levels is the single biggest challenge to implementing this approach. Donors agreed to help the government develop an Emergency Human Resources Programme with five main facets: improving incentives for recruitment and retention of staff through salary top-ups, expanding domestic training capacity, using international volunteer doctors and nurse tutors as a stop-gap measure, providing international technical assistance to bolster planning and management capacity and skills, and establishing more robust monitoring and evaluation capacity. Industrial relations were a prominent consideration in determining the shape of the Programme. The combination of short- and long-term measures appears to be helpful in maintaining commitment to the programme.
The purpose of this researcher was to examine the in-service experiences of seven headteachers in Luwinga ward in Mzuzu, Malawi, and the challenges they encountered in their first year on the job. A modified version of the questionnaire and interview guide developed by the members of the International Study of the Preparation of Principals at the…
Samu, Samu M.
This Peace Corps guide to individual, independent, or group study of Chichewa provides basic conversational vocabulary and phrases necessary for Peace Corps volunteer survival skills in Malawi. The 20 lessons consist of dialogues, vocabulary, and exercises on these topics: greetings; self-introduction; discussing others; relatives; professions;…
Chowa, Clodina; Garforth, Chris; Cardey, Sarah
Purpose: Malawi's current extension policy supports pluralism and advocates responsiveness to farmer demand. We investigate whether smallholder farmers' experience supports the assumption that access to multiple service providers leads to extension and advisory services that respond to the needs of farmers. Design/methodology/approach: Within a…
Baldauf, S A; Kullmann, H; Thünken, T; Winter, S; Bakker, T C M
Four choice experiments were conducted with both sexes of the cichlid Pelvicachromis taeniatus using computer-manipulated stimuli of digital images differing in movement, body shape or colouration. The results show that computer animations can be useful and flexible tools in studying preferences of a cichlid with complex and variable preferences for different visual cues.
Brown, E; Gorman, D; Knowles, G; Taylor, F; Jere, Y; Bates, J; Masamba, L
In response to the growing incidence of cancer in Malawi, a new oncology unit was established at the Queen Elizabeth Central Hospital, Blantyre. The unit opened in 2010, the first in the country, and is led by a single consultant oncologist. In 2012, a healthcare partnership was formed between the oncology and palliative care unit at Queen Elizabeth Central Hospital and the Edinburgh Cancer Centre, UK. The principal objective of the partnership is to help develop high quality multidisciplinary cancer care in Malawi. Methods A needs assessment identified three priority areas for further improvement of cancer services: nurse-led treatment delivery; management of clinical data; and multidisciplinary working. The partnership received grant funding from the Scottish Government Malawi Development Programme in 2013 and a three year project plan was implemented. This has been conducted through a series of reciprocal training visits. Results Key achievements have been completion of a programme of oncology nursing education attended by 32 oncology nurses and other healthcare professionals, which has resulted in increased experience in cancer practice and standardisation of chemotherapy delivery procedures; development of a clinical database that enables prospective collection of data of all new patients with cancer and which links to the Malawi Cancer Registry; development of weekly multidisciplinary meetings involving oncology, gynaecology and surgery that has enabled a cross-specialty approach to patient care. Conclusion The Edinburgh Malawi Cancer Partnership is supporting nursing education, data use and cross-specialty collaboration that we are confident will improve cancer care in Malawi. Future work will focus on the further development of multidisciplinary breast cancer care and the development of a radiotherapy service for patients in Malawi.
Bezault, Etienne; Rognon, Xavier; Gharbi, Karim; Baroiller, Jean-Francois; Chevassus, Bernard
The transfer of the genomic resources developed in the Nile tilapia, Oreochromis niloticus, to other Tilapiines sensu lato and African cichlid would provide new possibilities to study this amazing group from genetics, ecology, evolution, aquaculture, and conservation point of view. We tested the cross-species amplification of 32 O. niloticus microsatellite markers in a panel of 15 species from 5 different African cichlid tribes: Oreochromines (Oreochromis, Sarotherodon), Boreotilapiines (Tilapia), Chromidotilapines, Hemichromines, and Haplochromines. Amplification was successfully observed for 29 markers (91%), with a frequency of polymorphic (P95) loci per species around 70%. The mean number of alleles per locus and species was 3.2 but varied from 3.7 within Oreochromis species to 1.6 within the nontilapia species. The high level of cross-species amplification and polymorphism of the microsatellite markers tested in this study provides powerful tools for a wide range of molecular genetic studies within tilapia species as well as for other African cichlids. PMID:22701809
Kordesch, E. G.; Park, L. E.
The diversity of terrestrial systems is estimated to be greater than in the marine realm. However no hard data yet exists to substantiate this claim. Ancient lacustrine deposits may preserve an exceptionally diverse fossil fauna and aid in determining continental faunal diversities. Fossils preserved in lake deposits, especially those with exceptional preservation (i.e. Konservat Lagerstaetten), may represent a dependable method for determining species diversity changes in the terrestrial environment because of their faunal completeness. Important Konservat Lagerstaetten, such as the Green River Formation (US) and Messel (Germany), both Eocene in age, are found in lake sediments and show a remarkable faunal diversity for both vertebrates and invertebrates. To date information from nearly 25 lake lagerstaetten derived from different types of lake basins from the Carboniferous to the Miocene have been collected and described. Carboniferous sites derive from the cyclothems of Midcontinent of the US while many Cenozoic sites have been described from North and South America as well as Europe and Australia. Asian sites contain fossils from the Mesozoic and Cenozoic. With this data, insight into the evolutionary processes associated with lake systems can be examined. Do lakes act as unique evolutionary crucibles in contrast to marine systems? The speciation of cichlid fishes in present-day African lakes appears to be very high and is attributed to the diversity of environments found in large rift lakes. Is this true of all ancient lakes or just large rift lakes? The longevity of a lake system may be an important factor in allowing speciation and evolutionary processes to occur; marine systems are limited only in the existence of environments as controlled by tectonics and sea level changes, on the order of tens of millions of years. Rift lakes are normally the longest lived in the millions of years. Perhaps there are only certain types of lakes in which speciation of
Nagl, S; Mayer, W E; Klein, J
Complementary DNA clones coding for the catalytic unit of the enzyme glucose-6-phosphatase (G6Pase) were obtained from Haplochromis nubilus and Haplochromis xenognathus, two cichlid fish species from Lake Victoria. The translated sequence of these two cDNAs identifies a polypeptide consisting of 352 amino acid residues and showing a 54.4% similarity to the human form of G6Pase. The amino acid sequences of the two fish species are identical. The comparison of the fish amino acid sequence with the corresponding sequences of rat, mouse, and human G6Pase revealed that the amino acid residues, which are involved in G6Pase catalysis in humans, are also conserved in fish G6Pase. Northern blot analysis showed that G6Pase is expressed at the same level in 6- and 10-day-old fish. A three base pair insertion/deletion polymorphism was found in the 3'-untranslated region of the fish G6Pase gene. The polymorphism will be a useful marker in a phylogenetic study of Lake Victoria cichlids.
Henning, Frederico; Machado-Schiaffino, Gonzalo; Baumgarten, Lukas; Meyer, Axel
Genes of major phenotypic effects and strong genetic correlations can facilitate adaptation, direct selective responses and potentially lead to phenotypic convergence. However, the preponderance of this type of genetic architecture in repeatedly-evolved adaptations remains unknown. Using hybrids between Haplochromis chilotes (thick-lipped) and H. nyererei (thin-lipped) we investigated the genetics underlying hypertrophied lips and elongated heads, traits that evolved repeatedly in cichlids. At least 25 loci of small-to-moderate and mainly additive effects were detected. Phenotypic variation in lip and head morphology was largely independent. Although several QTL overlapped for lip and head morphology traits, they were often of opposite effects. The distribution of effect signs suggests strong selection on lips. The fitness implications of several detected loci were demonstrated using a laboratory assay testing for the association between genotype and variation in foraging performance. The persistence of low fitness alleles in head morphology appears to be maintained through antagonistic pleiotropy/close linkage with positive-effect lip morphology alleles. Rather than being based on few major loci with strong positive genetic correlations, our results indicate that the evolution of the Lake Victoria thick-lipped ecomorph is the result of selection on numerous loci distributed throughout the genome. This article is protected by copyright. All rights reserved.
Fitzpatrick, John L; Montgomerie, Robert; Desjardins, Julie K; Stiver, Kelly A; Kolm, Niclas; Balshine, Sigal
Sperm competition, the contest among ejaculates from rival males to fertilize ova of a female, is a common and powerful evolutionary force influencing ejaculate traits. During competitive interactions between ejaculates, longer and faster spermatozoa are expected to have an edge; however, to date, there has been mixed support for this key prediction from sperm competition theory. Here, we use the spectacular radiation of cichlid fishes from Lake Tanganyika to examine sperm characteristics in 29 closely related species. We provide phylogenetically robust evidence that species experiencing greater levels of sperm competition have faster-swimming sperm. We also show that sperm competition selects for increases in the number, size, and longevity of spermatozoa in the ejaculate of a male, and, contrary to expectations from theory, we find no evidence of trade-offs among sperm traits in an interspecific analysis. Also, sperm swimming speed is positively correlated with sperm length among, but not within, species. These different responses to sperm competition at intra- and interspecific levels provide a simple, powerful explanation for equivocal results from previous studies. Using phylogenetic analyses, we also reconstructed the probable evolutionary route of trait evolution in this taxon, and show that, in response to increases in the magnitude of sperm competition, the evolution of sperm traits in this clade began with the evolution of faster (thus, more competitive) sperm.
Dijkstra, Peter D; Lindström, Jan; Metcalfe, Neil B; Hemelrijk, Charlotte K; Brendel, Mischa; Seehausen, Ole; Groothuis, Ton G G
A mechanism commonly suggested to explain the persistence of color polymorphisms in animals is negative frequency-dependent selection. It could result from a social dominance advantage to rare morphs. We tested for this in males of red and blue color morphs of the Lake Victoria cichlid, Pundamilia. Earlier work has shown that males preferentially attack the males of their own morph, while red males are more likely to win dyadic contests with blue males. In order to study the potential contribution of both factors to the morph co-existence, we manipulated the proportion of red and blue males in experimental assemblages and studied its effect on social dominance. We then tried to disentangle the effects of the own-morph attack bias and social dominance of red using simulations. In the experiment, we found that red males were indeed socially dominant to the blue ones, but only when rare. However, blue males were not socially dominant when rare. The simulation results suggest that an own-morph attack bias reduces the social dominance of red males when they are more abundant. Thus, there is no evidence of symmetric negative frequency-dependent selection acting on social dominance, suggesting that additional fitness costs to the red morph must explain their co-existence.
Fitzpatrick, J L; Desjardins, J K; Milligan, N; Montgomerie, R; Balshine, S
Theory predicts that males experiencing elevated levels of sperm competition will invest more in gonads and produce faster-swimming sperm. Although there is ample evidence in support of the first prediction, few studies have examined sperm swimming speed in relation to sperm competition. In this study, we tested these predictions from sperm competition theory by examining sperm characteristics in Telmatochromis vittatus, a small shell-brooding cichlid fish endemic to Lake Tanganyika. Males exhibit four different reproductive tactics: pirate, territorial, satellite, and sneaker. Pirate males temporarily displace all other competing males from a shell nest, whereas sneaker males always release sperm in the presence of territorial and satellite males. Due to the fact that sneakers spawn in the presence of another male, sneakers face the highest levels of sperm competition and pirates the lowest, whereas satellites and territorials experience intermediate levels. In accordance with predictions, sperm from sneakers swam faster than sperm from males adopting the other reproductive tactics, whereas sperm from pirates was slowest. Interestingly, we were unable to detect any variation in sperm tail length among these reproductive tactics. Thus, sperm competition appears to have influenced sperm energetics in this species without having any influence on sperm size.
Ota, Kazutaka; Awata, Satoshi; Morita, Masaya; Yokoyama, Ryota; Kohda, Masanori
To examine how territorial males counter reproductive parasites, we examined the paternity of broods guarded by territorial males using 5 microsatellite loci and factors that determine siring success in a wild population of the Lake Tanganyika cichlid Lamprologus lemairii. Females enter rock holes (nests) and spawn inside, and territorial males release milt over the nest openings. Sneakers attempt to dart into the nests, but territorial males often interrupt the attempt. The body size of territorial males (territorial defense ability) and the size of nest opening (the ability to prevent sneakers from nest intrusions) are predicted to be factors that affect paternity at the premating stage, whereas milt quality traits are factors that affect paternity at the postmating stage. Parentage analyses of 477 offspring revealed that most clutches have few or no cuckolders, and territorial males sired >80% of eggs in 7 of the 10 analyzed clutches. Larger territorial males that spawned in nests with narrower openings had greater siring success. In contrast, none of the milt traits affected the siring success. These suggest that territorial male L. lemairii adopt premating strategies whereby they effectively prevent reproductive parasitism.
Heg, Dik; Bachar, Zina; Brouwer, Lyanne; Taborsky, Michael
Environmental conditions are thought to be responsible for the extent and benefits of cooperative breeding in many animal societies, but experimental tests are scarce. We manipulated predator pressure in the cooperatively breeding cichlid Neolamprologus pulcher in Lake Tanganyika, where predators have been suggested to influence helper and breeder survival, helper dispersal and group reproductive success. We varied the type and intensity of predation by releasing medium, large, or no predators inside large underwater cages enclosing two or three group territories. Helper and breeder survival, helper dispersal and group reproductive success decreased from the control, to the medium- and large-predator treatments. These effects were modified by helper body size and the number of adults protecting the group from predators, supporting the 'group augmentation hypothesis'. Predators forced helpers to stay closer to, and spend more time inside, protective shelters. The results demonstrate the importance of predators for group living in this species, and support the 'ecological constraints hypothesis' of cooperative breeding, in the sense that subordinates stay at home rather than leave and breed independently under the risk of predation. PMID:15556889
Bunje, Paul ME; Barluenga, Marta; Meyer, Axel
Background Speciation often occurs in complex or uncertain temporal and spatial contexts. Processes such as reinforcement, allopatric divergence, and assortative mating can proceed at different rates and with different strengths as populations diverge. The Central American Midas cichlid fish species complex is an important case study for understanding the processes of speciation. Previous analyses have demonstrated that allopatric processes led to species formation among the lakes of Nicaragua as well as sympatric speciation that is occurring within at least one crater lake. However, since speciation is an ongoing process and sampling genetic diversity of such lineages can be biased by collection scheme or random factors, it is important to evaluate the robustness of conclusions drawn on individual time samples. Results In order to assess the validity and reliability of inferences based on different genetic samples, we have analyzed fish from several lakes in Nicaragua sampled at three different times over 16 years. In addition, this time series allows us to analyze the population genetic changes that have occurred between lakes, where allopatric speciation has operated, as well as between different species within lakes, some of which have originated by sympatric speciation. Focusing on commonly used genetic markers, we have analyzed both DNA sequences from the complete mitochondrial control region as well as nuclear DNA variation at ten microsatellite loci from these populations, sampled thrice in a 16 year time period, to develop a robust estimate of the population genetic history of these diversifying lineages. Conclusion The conclusions from previous work are well supported by our comprehensive analysis. In particular, we find that the genetic diversity of derived crater lake populations is lower than that of the source population regardless of when and how each population was sampled. Furthermore, changes in various estimates of genetic diversity within lakes
Hyder, Asma; Behrman, Jere R.
This study investigates economic activities and their determinants for women in households of rural Malawi, one of the poorest countries in Sub-Saharan Africa (SSA). Three waves of household panel data for years 2006, 2008 and 2010 from the Malawi Longitudinal Study of Families and Health (MLSFH) are used to examine female labor market participation. This study includes certain important characteristics of Malawian households i.e., polygamy, ethnicity, religion and self reported health status. The study reports five different labor market outcomes for women's participation i.e., agriculture related work, market & sales activities, cottage industry, other economic activities and out of the labor market. The study finds significant relationships between these five different labor market outcomes and age cohorts, ethnicity and marriage (monogamous or polygamous). PMID:26120564
Suárez-Morales, Eduardo; Paredes-Trujillo, Amelia; González-Solís, David
The cyclopoid copepod Neoergasilus japonicus ( Harada, 1930 ) is recorded from three endangered or threatened fish species from southeast Mexico: the tailbar cichlid Vieja hartwegi (Taylor and Miller, 1980); the Angostura cichlid V. breidohri (Werner and Stawikowski, 1987); and the sieve cichlid C. grammodes (Taylor and Miller, 1980). This ectoparasitic copepod is considered, together with most other members of Neoergasilus, an Eastern Asian form. N. japonicus is one of the most widespread parasitic Asian copepods, as it has rapidly invaded Europe and North America, including Mexico. We estimated the prevalence, mean abundance, and intensity of infection of N. japonicus in these cichlid teleosts; our data agree with previous works stating the high prevalence of this ectoparasite. This copepod has a wide range of hosts among freshwater fish taxa, but this is only the second published report from cichlids in the Neotropical region. The three cichlids surveyed, V. hartwegi, V. breidohri, and C. grammodes, are new hosts of this copepod. Its occurrence in Mexico is attributed to different events of introduction by human agency. This is the southernmost record of N. japonicus in continental America. It is a matter of concern that this copepod is parasitizing endangered or threatened endemic cichlids in the Neotropical region. Because its high infective efficiency and ability to shift hosts, this Asian parasite is expected to spread farther southwards into Central and South America.
Malawi is one of the poorest countries in the world (World Bank, 2014). Yet, our experiences working with rural schools suggest that the Malawi education system may be far ahead of many developed nations, including the United States, in terms of their practical and philosophical commitment to inclusive education for all children, including…
Kamwendo, Gregory H.
According to the Medical Council of Malawi, one of the conditions for a licence to be granted to an individual who wants to practise medicine in Malawi is the practitioner's ability to speak and write English fluently. This means that the expatriate medical practitioner is not required by law to demonstrate fluency in Chichewa (the national…
Appiah-Yeboah, Shirley Afua
The circumstances that characterize poor, rural communities in Malawi suggest that children's health-wealth gradient can vary from other settings. This dissertation begins with a description of the methods used to create a household wealth variable using assets data in the Malawi Longitudinal Study of Families and Health project. By using a fixed…
Chasukwa Mwalwenje, Yvonne; Chasukwa, Steria
Malawi is popularly known as the Warm Heart of Africa. Malawi has a total land area of 45,747 sq. miles. Of the total area, 80 % is covered by fresh water from Lake Malawi and other rivers. The country boasts that it holds large amounts of fresh water and has the third largest lake in Africa. Ironically, the number of households with reliable water access is low (Water Aid, 2014, UNDP, Human Development Report 2008). Regardless of signs of economic development, water security still remains a challenge in the Warm Heart of Africa. The problem with access to water prompted the Government of Malawi to introduce a National Water Policy in 1994 with the vision of 'Water and Sanitation for All'. Since then, other water access interventions have impacted on the ability to manage water sustainably. Over the past few years, more inclusive and diversified interventions have been put in place to reverse the situation. For instance, the government of Malawi has taken action to increase the number of water tanks in schools (UNICEF 2005). Several stakeholders' support the government in their role provides policy direction and coordinate management of the water sector. The outlined National Water and Sanitation Policy strategies includes promoting water conservation and catchment protection; incorporating local governments and communities in planning, development and management of water supplies and sanitation services; rehabilitating the existing infrastructure; creating an enabling environment for public-private partnerships in water supply and sanitation activities; undertaking rehabilitation and reduction of unaccounted-for-water of existing urban, peri-urban, as a priority; promoting economic incentives and opportunities to encourage the participation of small-scale water and sanitation service providers; and promoting water recycling and re-use. Despite of all these interventions, Malawi still continues to face significant challenges with issues of access and quality of
Otañez, Marty G; Mamudu, Hadii; Glantz, Stanton A
Objective To examine the influence of US‐based tobacco leaf‐buying companies, Universal Corporation and Alliance One International, on Malawi's economy and trade policy in 2000–6. Design Analyses of ethnographic data and tobacco industry documents. Results Universal Corporation and Alliance One International, through their subsidiary companies Limbe Leaf and Alliance One, respectively, in Malawi, control policy‐making advisory groups and operate a tobacco cartel to influence Malawi's economic and trade sectors. Limbe Leaf's corporate secretary and lawyer is a member of several policy‐making committees that advise the Malawi government on tobacco‐related trade policy. The corporate representative's presence prevents other committee members from taking positions against the tobacco industry and ensures government policy that advances industry interests to obtain low‐cost tobacco. The World Bank and Malawi's Anti‐corruption Bureau report allegations of collusion between Limbe Leaf and Alliance One over prices at tobacco markets. Allegations of collusion between Limbe Leaf and Alliance One prompted Malawi President Bingu Mutharika in 2006 to warn the companies to end non‐competitive practices or leave the country, but there was no meaningful follow‐up action. Findings from interviews with small‐scale tobacco traders in Malawi suggest that Universal and Alliance One International purchase smuggled raw tobacco from the neighbouring countries, Zambia and Mozambique, undermining growers' efforts to benefit from tobacco farming in Malawi. Conclusion These actions restrict competition, depress tobacco prices for Malawi's farmers and contribute to poverty in Malawi, while keeping the country dependent on tobacco growing. PMID:17652242
Colgan, P W; Salmon, A B
Changes in the levels of activities by parental female and male convict cichlids were studied over three successive reproductive episodes and within each episode. No evidence was found for behavioural maturation independent of experience. With successive episodes, young survived for longer times, nests were cleaner, males ate less and females more, and males became more aggressive while females focused more on the nest. Within an episode there were also marked variations in activities by each sex. Reproductive behaviour is therefore open in a number of respects both within and across episodes to environmental inputs.
Smith, Alan; García-Alonso, Javier; van Oosterhout, Cock
Compared with other phylogenetic groups, self-fertilization (selfing) is exceedingly rare in vertebrates and is known to occur only in one small clade of fishes. Here we report observing one F1-hybrid individual that developed into a functional hermaphrodite after crossing two closely-related sexually reproducing species of cichlids. Microsatellite alleles segregated consistent with selfing and Mendelian inheritance and we could rule out different modes of parthenogenesis including automixis. We discuss why selfing is not more commonly observed in vertebrates in nature, and the role of hybridization in the evolution of novel traits. PMID:27069660
Maan, Martine E.; Sefc, Kristina M.
Cichlid fishes constitute one of the most species-rich families of vertebrates. In addition to complex social behaviour and morphological versatility, they are characterised by extensive diversity in colouration, both within and between species. Here, we review the cellular and molecular mechanisms underlying colour variation in this group and the selective pressures responsible for the observed variation. We specifically address the evidence for the hypothesis that divergence in colouration is associated with the evolution of reproductive isolation between lineages. While we conclude that cichlid colours are excellent models for understanding the role of animal communication in species divergence, we also identify taxonomic and methodological biases in the current research effort. We suggest that the integration of genomic approaches with ecological and behavioural studies, across the entire cichlid family and beyond it, will contribute to the utility of the cichlid model system for understanding the evolution of biological diversity. PMID:23665150
Hill, Jeffrey E.; Nico, Leo G.; Cichra, Charles E.; Gilbert, Carter R.
The interaction of prey fish body depth and predator gape size may produce prey assemblages dominated by invulnerable prey and excessive prey-to-predator biomass ratios. Peacock cichlids (Cichla ocellaris) were stocked into southeast Florida canals to consume excess prey fish biomass, particularly spotted tilapia (Tilapia mariae). The ecomorphologically similar largemouth bass (Micropterus salmoides) was already present in the canals. We present relations of length-specific gape size for peacock cichlids and largemouth bass. Both predators have broadly overlapping gape size, but largemouth bass ?126 mm total length have slightly larger gape sizes than peacock cichlids of the same length. Also, we experimentally tested the predictions of maximum prey size for peacock cichlids and determined that a simple method of measuring gape size used for largemouth bass also is appropriate for peacock cichlids. Lastly, we determined relations of body depth and length of prey species to investigate relative vulnerability. Using a simple predator-prey model and length frequencies of predators and bluegill (Lepomis macrochirus), redear sunfish (Lepomis microlophus), and spotted tilapia prey, we documented that much of the prey biomass in southeast Florida canals is unavailable for largemouth bass and peacock cichlid predation.
Zoppoth, P; Koblmüller, S; Sefc, K M
Whether premating isolation is achieved by male-specific, female-specific or sex-independent assortative preferences often depends on the underlying evolutionary processes. Here we test mate preferences of males presented with females of different allopatric colour variants of the cichlid fish Tropheus sp., a Lake Tanganyika endemic with rich geographical colour pattern variation, in which the strength of sexual isolation varies between populations. We conducted two-way mate choice experiments to compare behaviour of males of a red-bodied morph (population Moliro) towards females from their own population with behaviour towards females from four allopatric populations at different stages of phylogenetic and phenotypic divergence. Males courted same-population females significantly more intensely than females of other populations, and reduced their heteromorphic courtship efforts both with increasing genetic and increasing phenotypic distinctness of the females. In particular, females of a closely related red-bodied population received significantly more courtship than either genetically distinct, similarly coloured females ('Kirschfleck' morph) or genetically related, differently coloured females ('yellow-blotch' morph), both of which were courted similarly. Genetically and phenotypically distinct females (Tropheus polli) were not courted at all. Consistent with previous female-choice experiments, female courtship activity also decreased with increasing genetic distance from the males' population. Given successful experimental and natural introgression between colour morphs and the pervasive allopatry of related variants, we consider it unlikely that assortative preferences of both sexes were driven by direct selection during periods of secondary contact or, in turn, drove colour pattern differentiation in allopatry. Rather, we suggest that sexual isolation evolved as by-product of allopatric divergence.
Sengupta, Mita E; Kristensen, Thomas K; Madsen, Henry; Jørgensen, Aslak
The freshwater gastropod family Viviparidae is nearly cosmopolitan, but absent from South America. On the African continent, two genera are recognized; the widespread Bellamya and the monotypic Neothauma, which is confined to Lake Tanganyika. Most of the African Bellamya species are confined to the major lakes of the Rift Valley area in Africa, i.e. Lake Albert, Lake Malawi, Lake Mweru, and Lake Victoria. The phylogenetic analyses of mitochondrial (COI and 16S) and nuclear (H3, 18S and 28S) DNA inferred three major lake-clades; i.e. Lake Victoria/Kyoga/Albert, Lake Malawi and Lake Mweru/Bangweulu. The endemic B. rubicunda from Lake Albert and B. unicolor from Lake Kyoga were inferred to be part of the Lake Victoria clade. Bellamya capillata as identified by shell characters was polyphyletic in gene trees. The monophyletic Bellamya species radiation in Lake Malawi was most nearly related to the Lake Victoria/Kyoga/Albert-clade. Taxa from the Zambian lakes, Mweru and Bangweulu, were inferred together and placed ancestral to the other lakes. Neothauma tanganyicense was inferred as the sister-group to the Zambian Bellamya. Within the lake-clades the endemic radiations show very low genetic diversities (0-4.1% in COI), suggesting much faster morphological divergence than molecular divergence. Alternatively, Bellamya in Africa constitutes only a few species with several sub-species or eco-phenotypic morphs. The African viviparids were inferred to be the sister-group to a clade comprising Asian species, and the relatively low genetic diversity between the clades (12.6-15.5% in COI) makes a recent Miocene dispersal event from Asia to Africa much more likely than an ancient Gondwana vicarience distribution.
Hesse, Saskia; Anaya-Rojas, Jaime M; Frommen, Joachim G; Thünken, Timo
Kin selection theory predicts that cooperation is facilitated between genetic relatives, as by cooperating with kin an individual might increase its inclusive fitness. Although numerous theoretical papers support Hamilton's inclusive fitness theory, experimental evidence is still underrepresented, in particular in noncooperative breeders. Cooperative predator inspection is one of the most intriguing antipredator strategies, as it implies high costs on inspectors. During an inspection event, one or more individuals leave the safety of a group and approach a potential predator to gather information about the current predation risk. We investigated the effect of genetic relatedness on cooperative predator inspection in juveniles of the cichlid fish Pelvicachromis taeniatus, a species in which juveniles live in shoals under natural conditions. We show that relatedness significantly influenced predator inspection behaviour with kin dyads being significantly more cooperative. Thus, our results indicate a higher disposition for cooperative antipredator behaviour among kin as predicted by kin selection theory.
51 women with urinary schistosomiasis underwent gynecological examinations, including colposcopy and photographic documentation of lesions, in a study conducted in 1994 at Mangochi Hospital near Lake Malawi. Microscopy of genital biopsies determined that 33 women had S. haematobium eggs in their cervix, vagina, and/or vulva. A significant correlation was identified between the size of genital lesion and the number of ova counted. Women with genital pathology had significantly more tumors in the vulva than women with schistosomiasis of the urinary tract only. Most of the observed pathology could be seen with the unaided eye. Even though the sample was very small, significantly more cases of genital schistosomiasis were found in women who had fewer children than desired and whose husbands had children with other women. Reasons for that observation were not investigated. The findings of this pilot study indicate that more research is needed to provide a better understanding of the implications of the disease upon marital and sexual lives.
Malawi has been ranked by the World Bank as one of the poorest countries in Africa. Malawi's only resources are its people and fertile soil, which comprises about 55% of land area. Environmental degradation and population growth conditions in Malawi were used to illustrate the model of environmental degradation linked to population pressure on land resources and government development strategies that favored large-scale agricultural farms. The result has been deforestation, overgrazing, overuse of land for subsistence, and increased population density. The argument was that population growth in some developing countries has been so rapid that environmental collapse is the result. The theoretical framework linking population growth, environment, and resources emphasized processes: 1) the precursor stage of underlying causes; 2) the problem phase with potential ecological and economic decline; and 3) consequences (environmental decline, reduction in food production systems, and reduction in standard of living). The precursors were identified as an agrarian society, lack of a population policy, and emphasis on large families. The problems were rapid population growth and immigration from Mozambique, which led to increased demand for trees for fuel and consequent deforestation, increased demand for arable land and consequent landlessness, increased investment in livestock and consequent overgrazing, and continued population momentum which was a financial burden to government and resulted in increased labor competition. The ecological consequences were soil erosion, degradation of vegetation, and water supply contamination and decline. Eventually, famines will occur and lead to disease, migration, deserted villages, urbanization, unemployment, ethnic conflicts, and political unrest. Population was estimated at 8.75 million in 1990, with exponential growth expected. Completed family size was 6.6 children per woman. Even replacement fertility would mean growth for 50 more
Miller, K; Zulu, E M; Watkins, S C
Previous efforts by demographers to describe and explain spousal differences in reporting about family planning behavior have focused on individual attributes that are assumed to be related to the practice of contraception. This study extends that research by documenting spousal disagreement on a range of issues--household items, livestock, children, and spousal communication about fertility, family planning, and AIDS. Using data from a 1998 study of 585 monogamous couples in rural Malawi, the analysis identifies a systematic gender component to reporting: For many of the survey questions considered, when spouses disagree, husbands are more likely to say "yes" and wives "no." The findings are interpreted in terms of gendered strategies in the interview process.
Harries, A. D.; Kwanjana, J. H.; Hargreaves, N. J.; Van Gorkom, J.; Salaniponi, F. M.
OBJECTIVE: To document resources for controlling tuberculosis (TB) in Malawi. METHODS: We performed a countrywide study of all 43 hospitals (3 central, 22 district and 18 mission) which register and treat patients with TB. To collect data for 1998 on the TB-related workload, diagnostic facilities, programme staff and treatment facilities, we used laboratory, radiographic and TB registers, conducted interviews and visited hospital facilities. FINDINGS: The data show that in 1998, 88,257 TB suspects/patients contributed approximately 230,000 sputum specimens for smear microscopy, 55,667 chest X-rays were performed and 23,285 patients were registered for TB treatment. There were 86 trained laboratory personnel, 44 radiographers and 83 TB programme staff. Of these, about 40% had periods of illness during 1998. Approximately 20% of the microscopes and X-ray machines were broken. Some 16% of the hospital beds were designated for TB patients in special wards, but even so, the occupancy of beds in TB wards exceeded 100%. Although stocks of anti-TB drugs were good, there was a shortage of full-time TB ward nurses and 50% of district hospitals conducted no TB ward rounds. In general, there was a shortage of facilities for managing associated HIV-related disease; central hospitals, in particular, were underresourced. CONCLUSION: Malawi needs better planning to utilize its manpower and should consider cross-training hospital personnel. The equipment needs regular maintenance, and more attention should be paid to HIV-related illness. The policies of decentralizing resources to the periphery and increasing diagnostic and case-holding resources for central hospitals should be continued. PMID:11357212
Itimu, Ambumulire Nellie; Kopetz, Patricia B.
The sub-Saharan country of Malawi is steeped in a robust educational reform movement to ensure "Education for All". Based upon country immersion experience and a review of the literature, this report highlights Malawi's nationwide support for its special needs education (SNE) campaign. First discussed is a brief overview of Malawi, its…
In order to improve the fishery of Lake Victoria, a large predatory fish, the Nile perch (Lates niloticus), was introduced into the lake to feed on the smaller but abundant haplochromines, thereby producing a larger fish for food consumption. Originally the lake had a very high fish species diversity, dominated by more than 300 species of haplochromine cichlids; however, they were too small to be exploited for food. It took about 20 years for the population of the Nile perch in the lake to increase significantly. Although increased fish production has benefited riparian communities by providing revenue, employment, and food, the introduction of the Nile perch has also resulted in extinction and has transformed the lake ecosystem. In response to the concern caused by the introduction of the Nile perch, the deteriorating water quality, and the need to sustain the lucrative fishery, the governments of Uganda, Kenya, and Tanzania have initiated the multisectoral Lake Victoria Environment Management Programme. Its goals are to control the loading of nutrients and contaminants into the lake, manage land use in the catchment area, manage the introduced fish species, conserve biodiversity, and control the spread of water hyacinths.
Beeching, S C
Dummy conspecifics were presented to isolated adults of the cichlid fish Astronotus ocellatus to investigate the functional organization of cichlid social behavior. Body size and 15 dummy-elicited activities were recorded during 15 min sessions and analyzed by principal components analysis (PCA) to reveal their temporal organization. Five principal components explained almost 80% of the variation in dummy-elicited behavior, and these five factors define functional groups for Nest-oriented and attack modal action patterns are not mutually inhibitory during this time frame, and biting does not appear to function exclusively during an attack on a conspecific. Comparison with previous studies of New and Old World cichlids suggests evolutionary conservation of the functional organization of social behavior.
Thünken, Timo; Bakker, Theo C. M.; Kullmann, Harald
The main function of the spermatozoon is the transfer of the male haploid genome during fertilisation. In animals in general and in fishes in particular, there is huge variation in sperm size. In fishes, sperm size ranges from 13 μm in Mugil cephlus to nearly 100 μm in the channel catfish Ictalurus punctatus. We examined intra-specific variation in sperm morphometry in the socially monogamous cichlid Pelvicachromis taeniatus using scanning electron microscopy. The mean total sperm length of nearly 70 μm was extraordinarily large for cichlids. Furthermore, within-male variation was remarkably high. To our knowledge, P. taeniatus produces the longest cichlid sperm ever documented. Several hypotheses concerning the adaptive significance of these results are presented.
Ota, Kazutaka; Hori, Michio; Kohda, Masanori
To determine whether the appearance of a reproductively parasitic tactic varies, and how this variation affects territorial males of the Lake Tanganyika cichlid fish Telmatochromis vittatus, we examined the reproductive ecology of territorial males in Mtondwe and compared it with that of a neighboring Wonzye population, where nest density differs from that at Mtondwe. In Wonzye, with high nest density, male tactics change with their body size from a territorial to a non-territorial parasitic tactic called piracy in which they conquer several nests defended by territorial males and take over the nests while females are spawning. These "pirate" males could decrease the costs incurred by travelling among nests by exclusively targeting aggregations of nests in close proximity while avoiding separate nests. Territorial males in Wonzye sacrifice the potential higher attractiveness offered by large nests and instead compete for nests farther from neighbors on which pirates less frequently intrude. In contrast, the Mtondwe population had lower nest density and piracy was absent. Given that the success of piracy depends on the close proximity of nests, nest density is likely responsible for the observed variation in the occurrence of piracy between the two populations. Furthermore, in Mtondwe, territorial males competed for larger nests and were smaller than the territorial males in Wonzye. Thus, this lower nest density may free territorial males from the selection pressures for increased size caused by both defense against nest piracy and the need to develop into pirates as they grow.
Ota, Kazutaka; Hori, Michio; Kohda, Masanori
To determine whether the appearance of a reproductively parasitic tactic varies, and how this variation affects territorial males of the Lake Tanganyika cichlid fish Telmatochromis vittatus, we examined the reproductive ecology of territorial males in Mtondwe and compared it with that of a neighboring Wonzye population, where nest density differs from that at Mtondwe. In Wonzye, with high nest density, male tactics change with their body size from a territorial to a non-territorial parasitic tactic called piracy in which they conquer several nests defended by territorial males and take over the nests while females are spawning. These "pirate" males could decrease the costs incurred by travelling among nests by exclusively targeting aggregations of nests in close proximity while avoiding separate nests. Territorial males in Wonzye sacrifice the potential higher attractiveness offered by large nests and instead compete for nests farther from neighbors on which pirates less frequently intrude. In contrast, the Mtondwe population had lower nest density and piracy was absent. Given that the success of piracy depends on the close proximity of nests, nest density is likely responsible for the observed variation in the occurrence of piracy between the two populations. Furthermore, in Mtondwe, territorial males competed for larger nests and were smaller than the territorial males in Wonzye. Thus, this lower nest density may free territorial males from the selection pressures for increased size caused by both defense against nest piracy and the need to develop into pirates as they grow.
Berke, M. A.; Johnson, T. C.; Werne, J. P.; Schouten, S.; Sinninghe Damsté, J. S.
We present preliminary results from a study using the TEX86 temperature proxy from sediments of East African Rift Lakes (including Lakes Turkana, Albert, and Malawi) to reconstruct the thermal history of tropical Africa for the last ~ 20,000 years at a subcentennial to multicentennial resolution. The TEX86 proxy, based on tetraether membrane lipids produced by lacustrine Crenarchaeota, has been shown to be successful at recording lake surface temperatures of some large lakes, including Lakes Malawi and Tanganyika, while providing unreasonable surface temperatures for lakes that receive a large input of soil material. The East African Rift Lakes are climatically sensitive, with the majority of water loss due to evaporation rather than outflow. Thus, they are useful for paleoclimate studies, being sensitive to even small changes in aridity. Temperature records from the northern and central basins of Lake Malawi agree well and fall within modern surface lake temperatures. A 2.5°C cooling is evident during the Younger Dryas in the northern basin record, with no response seen in the central basin. We are currently investigating mechanisms to explain why both records show a gradual cooling of 3°C during the late Holocene. Lake Albert shows an intriguing two-step cooling during the Younger Dryas, reaching temperatures 2.5°C lower than temperatures preceding or following this interval. The temperature record of Lake Turkana shows an interesting ~ 500 year cyclicity of low temperatures punctuated by abrupt warming events. Lakes Turkana and Albert show TEX86 paleotemperatures considerably lower (8°C cooler in Lake Albert and ~ 4°C cooler in Lake Turkana) than modern surface water temperatures. Although these records appear to fall in the range of temporal variability, these temperature discrepancies may indicate varying Crenarcheotal populations between lakes or other influencing factors.
Nichols, Paul; Genner, Martin J; van Oosterhout, Cock; Smith, Alan; Parsons, Paul; Sungani, Harold; Swanstrom, Jennifer; Joyce, Domino A
Theory proposes that genomic admixture between formerly reproductively isolated populations can generate phenotypic novelty for selection to act upon. Secondary contact may therefore be a significant promoter of phenotypic novelty that allows species to overcome environmental challenges and adapt to novel environments, including during adaptive radiation. To date, this has largely been considered from the perspective of interspecific hybridization at contact zones. However, it is also possible that this process occurs more commonly between natural populations of a single species, and thus its importance in adaptive evolution may have been underestimated. In this study, we tested the consequences of genomic introgression during apparent secondary contact between phenotypically similar lineages of the riverine cichlid fish Astatotilapia calliptera. We provide population genetic evidence of a secondary contact zone in the wild, and then demonstrate using mate-choice experiments that both lineages can reproduce together successfully in laboratory conditions. Finally, we show that genomically admixed individuals display extreme phenotypes not observed in the parental lineages. Collectively, the evidence shows that secondary contact can drive the evolution of phenotypic novelty, suggesting that pulses of secondary contact may repeatedly seed genetic novelty, which when coupled with ecological opportunity could promote rapid adaptive evolution in natural circumstances.
Maruska, Karen P.; Fernald, Russell D.
SUMMARY Chemosensory signaling is crucial for communication in many fish species, but little is known about how signalers modulate chemical output in response to sensory information and social context. Here, we tested the hypothesis that dominant male African cichlid fish (Astatotilapia burtoni) use urine signals during social interactions, and demonstrate that this signaling depends on social context (reproductive; territorial) and on available sensory information (visual cues; full interaction). We injected males with dye to visualize urine pulses and exposed them to full sensory information or visual cues alone of four types: (1) dominant male; (2) gravid (reproductively receptive) females; (3) mouth-brooding (non-receptive) females; or (4) control (no fish). We found that males released urine sooner and increased their urination frequency when visually exposed to gravid females as compared with mouth-brooding females and or no-fish controls. While males could distinguish female reproductive states using visual cues alone, courtship behavior rates were ∼10-fold higher when they fully interacted with gravid females compared with receiving visual cues alone. Males also increased their urination and territorial behaviors when exposed to another male, suggesting that chemical signals may convey information on dominance status. These data support the hypothesis that dominant males use urine as a chemical signal and adjust the frequency of their urine output based on contextual information. PMID:22162854
Langen, Kathrin; Schwarzer, Julia; Kullmann, Harald; Bakker, Theo C M; Thünken, Timo
In wild animal populations, the degree of inbreeding differs between species and within species between populations. Because mating with kin often results in inbreeding depression, observed inbreeding is usually regarded to be caused by limited outbreeding opportunities due to demographic factors like small population size or population substructuring. However, theory predicts inclusive benefits from mating with kin, and thus part of the observed variation in inbreeding might be due to active inbreeding preferences. Although some recent studies indeed report kin mating preferences, the evidence is still highly ambiguous. Here, we investigate inbreeding in a natural population of the West African cichlid fish Pelvicachromis taeniatus which showed clear kin mating preferences in standardized laboratory experiments but no inbreeding depression. The presented microsatellite analysis reveals that the natural population has, in comparison to two reference populations, a reduced allelic diversity (A = 3) resulting in a low heterozygosity (H(o) = 0.167) pointing to a highly inbred population. Furthermore, we found a significant heterozygote deficit not only at population (F(is) = 0.116) but also at subpopulation level (F(is) = 0.081) suggesting that inbreeding is not only a by-product of population substructuring but possibly a consequence of behavioral kin preferences.
Nichols, Paul; Genner, Martin J.; van Oosterhout, Cock; Smith, Alan; Parsons, Paul; Sungani, Harold; Swanstrom, Jennifer; Joyce, Domino A.
Theory proposes that genomic admixture between formerly reproductively isolated populations can generate phenotypic novelty for selection to act upon. Secondary contact may therefore be a significant promoter of phenotypic novelty that allows species to overcome environmental challenges and adapt to novel environments, including during adaptive radiation. To date, this has largely been considered from the perspective of interspecific hybridization at contact zones. However, it is also possible that this process occurs more commonly between natural populations of a single species, and thus its importance in adaptive evolution may have been underestimated. In this study, we tested the consequences of genomic introgression during apparent secondary contact between phenotypically similar lineages of the riverine cichlid fish Astatotilapia calliptera. We provide population genetic evidence of a secondary contact zone in the wild, and then demonstrate using mate-choice experiments that both lineages can reproduce together successfully in laboratory conditions. Finally, we show that genomically admixed individuals display extreme phenotypes not observed in the parental lineages. Collectively, the evidence shows that secondary contact can drive the evolution of phenotypic novelty, suggesting that pulses of secondary contact may repeatedly seed genetic novelty, which when coupled with ecological opportunity could promote rapid adaptive evolution in natural circumstances. PMID:25392475
Galhardo, L.; Vitorino, A.; Oliveira, R. F.
Personality traits, such as exploration–avoidance, are expected to be adaptive in a given context (e.g. low-risk environment) but to be maladaptive in others (e.g. high-risk environment). Therefore, it is expected that personality traits are flexible and respond to environmental fluctuations, given that consistency across different contexts is maintained, so that the relative individual responses in relation to others remains the same (i.e. although the magnitude of the response varies the differences between high and low responders are kept). Here, we tested the response of male cichlid fish (Oreochromis mossambicus) to a novel object (NO) in three different social contexts: (i) social isolation, (ii) in the presence of an unfamiliar conspecific, and (iii) in the presence of a familiar conspecific. Males in the familiar treatment exhibited more exploratory behaviour and less neophobia than males in either the unfamiliar or the social isolation treatments. However, there were no overall correlations in individual behaviour across the three treatments, suggesting a lack of consistency in exploration–avoidance as measured by the NO test in this species. Moreover, there were no differences in cortisol responsiveness to an acute stressor between the three treatments. Together, these results illustrate how behavioural traits usually taken as measures of personality may exhibit significant flexibility and lack the expected consistency across different social contexts. PMID:22859562
Huffman, Lin S; O'Connell, Lauren A; Hofmann, Hans A
The roles of estrogen and androgens in male social behavior are well studied, but little is known about how these hormones contribute to behavior in a social hierarchy. Here we test the role of aromatase, the enzyme that converts testosterone into estradiol, in mediating aggression and reproductive behavior in male Astatotilapia burtoni, an African cichlid fish that displays remarkable plasticity in social behavior. We first measured aromatase expression in subordinate and dominant males in brain regions that regulate social behavior and found that subordinate males have higher aromatase expression than dominant males in the magnocellular and gigantocellular regions of the preoptic area. Next, we functionally tested the role of aromatase in regulating behavior by intraperitoneally injecting dominant males with either saline or fadrozole (FAD), an aromatase inhibitor, and found that FAD treatment decreases aggressive, but not reproductive, behaviors compared to saline controls. To determine the underlying physiological and molecular consequences of FAD treatment, we measured estradiol and testosterone levels from plasma and brain aromatase expression in FAD and saline treated dominant males. We found that estradiol levels decreased and testosterone levels increased in response to FAD treatment. Moreover, FAD treated males had increased aromatase expression in the gigantocellular portion of the POA, possibly a compensatory response. Overall, our results suggest aromatase is a key enzyme that promotes aggression in A. burtoni males through actions in the preoptic area.
Galhardo, L; Vitorino, A; Oliveira, R F
Personality traits, such as exploration-avoidance, are expected to be adaptive in a given context (e.g. low-risk environment) but to be maladaptive in others (e.g. high-risk environment). Therefore, it is expected that personality traits are flexible and respond to environmental fluctuations, given that consistency across different contexts is maintained, so that the relative individual responses in relation to others remains the same (i.e. although the magnitude of the response varies the differences between high and low responders are kept). Here, we tested the response of male cichlid fish (Oreochromis mossambicus) to a novel object (NO) in three different social contexts: (i) social isolation, (ii) in the presence of an unfamiliar conspecific, and (iii) in the presence of a familiar conspecific. Males in the familiar treatment exhibited more exploratory behaviour and less neophobia than males in either the unfamiliar or the social isolation treatments. However, there were no overall correlations in individual behaviour across the three treatments, suggesting a lack of consistency in exploration-avoidance as measured by the NO test in this species. Moreover, there were no differences in cortisol responsiveness to an acute stressor between the three treatments. Together, these results illustrate how behavioural traits usually taken as measures of personality may exhibit significant flexibility and lack the expected consistency across different social contexts.
Van Wassenbergh, Sam; Potes, Nuno Zavattieri; Adriaens, Dominique
Presumably as an adaptation for mouthbrooding, many cichlid fish species have evolved a prominent sexual dimorphism in the adult head. Since the head of fishes serves as a bow during locomotion, an evolutionary increase in head volume to brood more eggs can trade-off with the hydrodynamic efficiency of swimming. Here, the differences between males and females in three-dimensional shape and size of the external head surfaces and the effect thereof on drag force during locomotion was analysed for the Nile tilapia (Oreochromis niloticus), a maternal mouthbrooder. To do so, three-dimensional body surface reconstructions from laser scans and computational fluid dynamics simulations were performed. After scaling the scanned specimens to post-cranial body volume, in order to theoretically equalize propulsive power, the external volume of the head of females was 27% larger than that of males (head length + 14%; head width + 9%). These differences resulted in an approximate 15% increase in drag force. Yet, hydrodynamics imposed important constraints on the adaptation for mouthbrooding as a much more drastic drop in swimming efficiency seems avoided by mainly enlarging the head along the swimming direction.
Hesse, Saskia; Anaya-Rojas, Jaime M.; Frommen, Joachim G.; Thünken, Timo
The social environment individuals are exposed to during ontogeny shapes social skills and social competence in group-living animals. Consequently, social deprivation has serious effects on behaviour and development in animals but little is known about its impact on cooperation. In this study, we examined the effect of social environment on cooperative predator inspection. Predator inspection behaviour is a complex behaviour, which is present in a variety of shoaling fish species. Often, two fish leave the safety of the group and inspect a potentially dangerous predator in order to gather information about the current predation risk. As predator inspection is highly risky, it is prone to conflicts and cheating. However, cooperation among individuals may reduce the individual predation risk. We investigated this complex social behaviour in juveniles of the cichlid fish Pelvicachromis taeniatus that were reared in two different social environments throughout development. Fish reared in a group inspected more often than isolation-reared fish and were more likely to cooperate, i.e. they conducted conjoint inspection of a predator. By contrast, isolation-reared fish were more likely to perform a single inspection without a companion. These results suggest an impairment of cooperative behaviour in isolation-reared fish most probably due to lack of social experience and resulting in lowered social skills needed in coordinated behaviour. PMID:26064616
Antunes, Raquel A.; Oliveira, Rui F.
In many territorial species androgens respond to social interactions. This response has been interpreted as a mechanism for adjusting aggressive motivation to a changing social environment. Therefore, it would be adaptive to anticipate social challenges and reacting to their clues with an anticipatory androgen response to adjust agonistic motivation to an imminent social challenge. Here we test the hypothesis of an anticipatory androgen response to territorial intrusions using classical conditioning to establish an association between a conditioned stimulus (CS = light) and an unconditioned stimulus (US = intruder male) in male cichlid fish (Oreochromis mossambicus). During the training phase conditioned males (CS−US paired presentations) showed a higher decrease in latency for agonistic response toward the intruder than unconditioned males (CS–US unpaired presentations). In the test trial, conditioned males showed an increase in androgen levels (i.e., testosterone and 11-ketotestosterone) relative to baseline, in response to the CS alone. This increase was similar to that of control males exposed to real intruders after CS, whereas unconditioned males showed a decrease in androgen levels in response to the CS. Furthermore, conditioned males were significantly more aggressive than unconditioned males during the post-CS period on test trial, even though the intruder male was not present during this period. These results reveal the occurrence of a conditioned androgen response that may give territorial males an advantage in mounting a defense to upcoming territorial intrusions, if the ability to readily elevate androgens does not co-vary with other traits that bear costs. PMID:19805238
Nichols, E K; Giles, D; Kang'oma, S; Mwalwanda, L; Onaka, A; Notzon, F
In 2010, Malawi adopted a National Registration Act, making the registration of births and deaths compulsory, and efforts to improve Malawi's civil registration and vital statistics (CRVS) system are underway. During a participatory-style workshop, stakeholders completed a rapid assessment of the national civil registration and vital statistics systems. While participants discussed and scored each item in a standard tool, the workshop focused on sharing of partners' roles and challenges. The workshop has enhanced receptiveness in collaboration, and an inter-ministerial technical working group has now been formed to develop a strategic plan and conduct a comprehensive assessment to guide future improvements.
Giles, D.; Kang'oma, S.; Mwalwanda, L.; Onaka, A.; Notzon, F.
In 2010, Malawi adopted a National Registration Act, making the registration of births and deaths compulsory, and efforts to improve Malawi's civil registration and vital statistics (CRVS) system are underway. During a participatory-style workshop, stakeholders completed a rapid assessment of the national civil registration and vital statistics systems. While participants discussed and scored each item in a standard tool, the workshop focused on sharing of partners' roles and challenges. The workshop has enhanced receptiveness in collaboration, and an inter-ministerial technical working group has now been formed to develop a strategic plan and conduct a comprehensive assessment to guide future improvements. PMID:26399284
Maleta, K; Virtanen, S; Espo, M; Kulmala, T; Ashorn, P
Aim: To determine the timing of growth faltering among under 3 year old children. Methods: Prospective population based cohort study in Lungwena, rural Malawi, southeast Africa. A total of 767 live born babies were regularly visited from birth until 3 years of age. Weight, height, and mid upper arm circumference were measured at monthly intervals until 18 months and at three month intervals thereafter. Growth charts were constructed using the LMS method and comparisons made to two international databases: the traditional United States National Center for Health Statistics/World Health Organisation (NCHS/WHO) reference and the recently developed 2000 Centers for Disease Control (CDC) growth reference. Results: Compared to the 2000 CDC reference population, newborns in Lungwena were on average 2.5 cm shorter and 510 g lighter. On a population level, height faltering was present at birth and continued throughout the first three years. Weight faltering, on the other hand, occurred mainly between 3 and 12 months of age. At 36 months, the mean weight and height of the study children were 2.3 kg and 10.5 cm lower than those of the reference population, respectively. The results remained essentially similar when the comparisons were made to the NCHS/WHO reference. Conclusions: The fact that weight and height faltering do not follow identical time patterns suggests that they may have different origin and determinants. Further studies on the aetiology of height faltering and different approaches to preventive interventions are needed. PMID:12818899
Lowe, R.; Chadza, T.; Chirombo, J.; Fonda, C.; Muyepa, A.; Nkoloma, M.; Pietrosemoli, E.; Radicella, S. M.; Tompkins, A. M.; Zennaro, M.
It is commonly accepted that climate plays a role in the transmission of many infectious diseases, particularly those transmitted by mosquitoes such as malaria, which is one of the most important causes of mortality and morbidity in developing countries. Due to time lags involved in the climate-disease transmission system, lagged observed climate variables could provide some predictive lead for forecasting disease epidemics. This lead time could be extended by using forecasts of the climate in disease prediction models. This project aims to implement a platform for the dissemination of climate-driven disease risk forecasts, using a telemedicine approach. A pilot project has been established in Malawi, where a 162 km wireless link has been installed, spanning from Blantyre City to remote health facilities in the district of Mangochi in the Southern region, bordering Lake Malawi. This long Wi-Fi technology allows rural health facilities to upload real-time disease cases as they occur to an online health information system (DHIS2); a national medical database repository administered by the Ministry of Health. This technology provides a real-time data logging system for disease incidence monitoring and facilitates the flow of information between local and national levels. This platform allows statistical and dynamical disease prediction models to be rapidly updated with real-time climate and epidemiological information. This permits health authorities to target timely interventions ahead of an imminent increase in malaria incidence. By integrating meteorological and health information systems in a statistical-dynamical prediction model, we show that a long-distance Wi-Fi link is a practical and inexpensive means to enable the rapid analysis of real-time information in order to target disease prevention and control measures and mobilise resources at the local level.
Sefc, Kristina M.
Fish color patterns are among the most diverse phenotypic traits found in the animal kingdom. Understanding the molecular and cellular mechanisms that control in chromatophore distribution and pigmentation underlying this diversity is a major goal in developmental and evolutionary biology, which has predominantly been pursued in the zebrafish model system. Here, we apply results from zebrafish work to study a naturally occurring color pattern phenotype in the fins of an African cichlid species from Lake Tanganyika. The cichlid fish Neolamprologus meeli displays a distinct dorsal color pattern, with black and white stripes along the edges of the dorsal fin and of the dorsal half of the caudal fin, corresponding with differences in melanophore density. To elucidate the molecular mechanisms controlling the differences in dorsal and ventral color patterning in the fins, we quantitatively assessed the expression of 15 candidate target genes involved in adult zebrafish pigmentation and stripe formation. For reference gene validation, we screened the expression stability of seven widely expressed genes across the investigated tissue samples and identified tbp as appropriate reference. Relative expression levels of the candidate target genes were compared between the dorsal, striped fin regions and the corresponding uniform, grey-colored regions in the anal and ventral caudal fin. Dorso-ventral expression differences, with elevated levels in both white and black stripes, were observed in two genes, the melanosome protein coding gene pmel and in igsf11, which affects melanophore adhesion, migration and survival. Next, we predicted potential shared upstream regulators of pmel and igsf11. Testing the expression patterns of six predicted transcriptions factors revealed dorso-ventral expression difference of irf1 and significant, negative expression correlation of irf1 with both pmel and igsf11. Based on these results, we propose pmel, igsf11 and irf1 as likely components of
Böhne, Astrid; Sengstag, Thierry; Salzburger, Walter
Males and females of the same species differ largely in gene expression, which accounts for most of the morphological and physiological differences and sex-specific phenotypes. Here, we analyzed sex-specific gene expression in the brain and the gonads of cichlid fishes from Lake Tanganyika belonging to four different lineages, so-called tribes (Eretmodini, Ectodini, Haplochromini, and Lamprologini), using the outgroup Nile tilapia (Oreochromis niloticus) as reference. The comparison between male and female brains revealed few differences between the sexes, consistent in all investigated species. The gonads, on the other hand, showed a large fraction of differentially expressed transcripts with the majority of them showing the same direction of expression in all four species. All here-studied cichlids, especially the three investigated mouth-breeding species, showed a trend toward more male- than female-biased transcripts. Transcripts, which were female-biased in expression in all four species, were overrepresented on linkage group (LG)1 in the reference genome and common male-biased transcripts showed accumulation on LG23, the presumable sex chromosomes of the Nile tilapia. Sex-specific transcripts contained candidate genes for sex determination and differentiation in fishes, especially members of the transforming growth factor-β-superfamily and the Wnt-pathway and also prominent members of the sox-, dm-domain-, and high mobility group-box families. We further confirmed our previous finding on species/lineage-specific gene expression shifts in the sex steroid pathway, including synthesizing enzymes as the aromatase cyp19a1 and estrogen and androgen receptors. PMID:25364805
Steele, Sarah E.; López-Fernández, Hernán
Body size is an important correlate of life history, ecology and distribution of species. Despite this, very little is known about body size evolution in fishes, particularly freshwater fishes of the Neotropics where species and body size diversity are relatively high. Phylogenetic history and body size data were used to explore body size frequency distributions in Neotropical cichlids, a broadly distributed and ecologically diverse group of fishes that is highly representative of body size diversity in Neotropical freshwater fishes. We test for divergence, phylogenetic autocorrelation and among-clade partitioning of body size space. Neotropical cichlids show low phylogenetic autocorrelation and divergence within and among taxonomic levels. Three distinct regions of body size space were identified from body size frequency distributions at various taxonomic levels corresponding to subclades of the most diverse tribe, Geophagini. These regions suggest that lineages may be evolving towards particular size optima that may be tied to specific ecological roles. The diversification of Geophagini appears to constrain the evolution of body size among other Neotropical cichlid lineages; non-Geophagini clades show lower species-richness in body size regions shared with Geophagini. Neotropical cichlid genera show less divergence and extreme body size than expected within and among tribes. Body size divergence among species may instead be present or linked to ecology at the community assembly scale. PMID:25180970
Cazzaniga, Ilaria; Giardino, Claudia; Bresciani, Mariano; Poser, Kathrin; Peters, Steef; Hommersom, Annelies; Schenk, Karin; Heege, Thomas; Philipson, Petra; Ruescas, Ana; Bottcher, Martin; Stelzer, Kerstin
The aim of this study is the analysis of trend in the trophic level evolution in clear deep lakes which, being characterised by good quality state, are important socio- economic resources for their regions. The selected lakes are situated in Europe (Garda, Maggiore, Constance and Vättern), North America (Michigan) and Africa (Malawi and Tanganyika) and cover a range of eco- regions (continental, perialpine, boreal, rift valley) distributed globally.To evaluate trophic level tendency we mainly focused on chlorophyll-a concentrations (chl-a) which is a direct proxy of trophic status. The chl-a concentrations were obtained from 5216 cloud-free MERIS imagery from 2002 to 2012.The 'GLaSS RoIStats tool' available within the GLaSS project was used to extract chl-a in a number of region of interests (ROI) located in pelagic waters as well as some few other stations depending on lakes morphology. For producing the time-series trend, these extracted data were analysed with the Seasonal Kendall test.The results overall show almost stable conditions with a slight increase in concentration for lakes Maggiore, Constance, and the Green Bay of Lake Michigan; a slight decrease for lakes Garda and Tanganyika and absolutely stable conditions for lakes Vättern and Malawi.The results presented in this work show the great capability of MERIS to perform trend tests analysis on trophic status with focus on chl-a concentration. Being chl-a also a key parameter in water quality monitoring plans, this study also supports the managing practices implemented worldwide for using the water of the lakes.
Background Transposable elements (TEs) have the potential to produce broad changes in the genomes of their hosts, acting as a type of evolutionary toolbox and generating a collection of new regulatory and coding sequences. Several TE classes have been studied in Neotropical cichlids; however, the information gained from these studies is restricted to the physical chromosome mapping, whereas the genetic diversity of the TEs remains unknown. Therefore, the genomic organization of the non-LTR retrotransposons Rex1, Rex3, and Rex6 in five Amazonian cichlid species was evaluated using physical chromosome mapping and DNA sequencing to provide information about the role of TEs in the evolution of cichlid genomes. Results Physical mapping revealed abundant TE clusters dispersed throughout the chromosomes. Furthermore, several species showed conspicuous clusters accumulation in the centromeric and terminal portions of the chromosomes. These TE chromosomal sites are associated with both heterochromatic and euchromatic regions. A higher number of Rex1 clusters were observed among the derived species. The Rex1 and Rex3 nucleotide sequences were more conserved in the basal species than in the derived species; however, this pattern was not observed in Rex6. In addition, it was possible to observe conserved blocks corresponding to the reverse transcriptase fragment of the Rex1 and Rex3 clones and to the endonuclease of Rex6. Conclusion Our data showed no congruence between the Bayesian trees generated for Rex1, Rex3 and Rex6 of cichlid species and phylogenetic hypothesis described for the group. Rex1 and Rex3 nucleotide sequences were more conserved in the basal species whereas Rex6 exhibited high substitution rates in both basal and derived species. The distribution of Rex elements in cichlid genomes suggests that such elements are under the action of evolutionary mechanisms that lead to their accumulation in particular chromosome regions, mostly in heterochromatins. PMID
Hulsey, C D; García de León, F J; Rodiles-Hernández, R
The extent to which elements of functional systems can change independently (modularity) likely influences the diversification of lineages. Major innovations in organismal design, like the pharyngeal jaw in cichlid fishes, may be key to a group's success when they relax constraints on diversification by increasing phenotypic modularity. In cichlid fishes, pharyngeal jaw modifications that enhanced the ability to breakdown prey may have freed their oral jaws from serving their ancestral dual role as a site of both prey capture and prey processing. This functional decoupling that allowed the oral jaws to become devoted solely to prey capture has been hypothesized to have permitted the two sets of cichlid jaws to evolve independently. We tested the hypothesis that oral and pharyngeal jaw mechanics are evolutionarily decoupled both within and among Neotropical Heroine cichlids. In the trophically polymorphic species Herichthys minckleyi, molariforms that exhibit enlarged molarlike pharyngeal jaw teeth were found to have approximately 400% greater lower jaw mass compared to H. minckleyi with the alternative papilliform pharyngeal morphology. However, oral jaw gape, lower jaw velocity ratios, anterior jaw linkage mechanics, and jaw protrusion did not differ between the morphotypes. In 40 other Heroine species, there was a weak correlation between oral jaw mechanics and pharyngeal jaw mass when phylogenetic history was ignored. Yet, after expansion of the cytochrome b phylogeny for Heroines, change in oral jaw mechanics was found to be independent of evolutionary change in pharyngeal jaw mass based on independent contrasts. Evolutionary decoupling of oral and pharyngeal jaw mechanics has likely played a critical role in the unparalleled trophic diversification of cichlid fishes.
This article presents an account of a practical outdoor science project using the larvae of antlions with year 5 and 6 (age 10-11) pupils from Sir Harry Johnston International Primary School in Zomba, Malawi. Aim: To conduct a science project outdoors using scientific enquiry and knowledge in a typical African school environment and to see whether…
Fedderke, Johannes; Garlick, Julia
This paper constructs a new set of institutional indicators for Malawi. We develop indicators of political rights, of freehold, traditional (communitarian) and intellectual property rights, based on the Malawian legislative framework. In exploring the association between our rights measures and a range of indicators of socio-economic development,…
Malawi was the first country after the Jomtien conference to offer free primary education. Despite this policy, universal education has remained elusive. Many children do enroll in school, but not at the official/recommended age and drop out before completing school. Understanding these transition points--entry and completion--is critical to the…
Grant, Monica; Lloyd, Cynthia; Mensch, Barbara
The provision of toilets and menstrual supplies appears to be a promising strategy to promote adolescent girls' school attendance and performance in less developed countries. In this article, we use the first round of the Malawi Schooling and Adolescent Survey (MSAS) to examine the individual- and school-level factors associated with…
Chimombo, Joseph Patrick Godson
Free primary education was introduced to Malawi in 1994, and it led to an increase in enrollment of one million pupils. The present contribution gathers case studies of 10 schools which sought information on the general conditions of communities and schools as well as on enrollment, resources, staffing, absenteeism etc. The results highlight the…
Tavory, Iddo; Swidler, Ann
This article examines the widespread resistance to condom use in sub-Saharan Africa by describing the major semiotic axes that organize how people talk about condoms and condom use. These axes include the "sweetness" of sex, trust and love between sexual partners, and assessments of risk and danger. Using data from rural Malawi, we show…
Harries, A. D.; Hargreaves, N. J.; Gausi, F.; Kwanjana, J. H.; Salaniponi, F. M.
OBJECTIVE: Following the introduction of guidelines for the control of tuberculosis (TB) infection in all hospitals in Malawi, a study was carried out to determine whether the guidelines were being implemented, the time between admission to hospital and the diagnosis of pulmonary TB had been reduced, and the annual case notification rates among health workers had fallen and were comparable to those of primary-school teachers. METHODS: The study involved 40 district and mission hospitals. Staff and patients were interviewed in order to determine whether the guidelines had been adopted. In four hospitals the diagnostic process in patients with smear-positive pulmonary TB was evaluated before and after the introduction of the guidelines, with the aid of case notes and TB registers. In all hospitals the proportion of health workers registered with TB before and after the guidelines were introduced, in 1996 and 1999, respectively, was determined by conducting interviews and consulting staff lists and TB registers. A similar method was used to determine the proportion of primary-school teachers who were registered with TB in 1999. FINDINGS: The guidelines were not uniformly implemented. Only one hospital introduced voluntary counselling and testing for its staff. Most hospitals stated that they used rapid systems to diagnose pulmonary TB. However, there was no significant change in the interval between admission and diagnosis or between admission and treatment of patients with smear-positive pulmonary TB. The TB case notification rate for 2979 health workers in 1999 was 3.2%; this did not differ significantly from the value of 3.7% for 2697 health workers in 1996 but was significantly higher than that of 1.8% for 4367 primary-school teachers in 1999. CONCLUSION: The introduction of guidelines for the control of TB infection is an important intervention for reducing nosocomial transmission of the disease, but rigorous monitoring and follow-up are needed in order to ensure
Chapman, L.J.; Chapman, Colin A.; Schofield, P.J.; Olowo, J.P.; Kaufman, L.S.; Seehausen, O.; Ogutu-Ohwayo., R.
In Lake Nabugabo, Uganda, a small satellite of the equatorial Lake Victoria, approximately 50% of the indigenous fish species disappeared from the open waters subsequent to establishment of the introduced predatory Nile perch ( Lates niloticus ). However, several of these species persisted in wetland refugia. Over the past decade, Nile perch in Lake Nabugabo have been intensively fished. Herein we report a resurgence of some indigenous species in open waters. In a multiyear study, we used annual transects in inshore and offshore waters of exposed ( no wetland ) and wetland habitats to document the pattern of resurgence. In 1995, haplochromine cichlids were largely confined to inshore areas, particularly wetland ecotones, and were rare in Nile perch stomachs, as were most other indigenous species. By 2000 haplochromine cichlids were abundant in inshore and offshore areas of both exposed and wetland transects. Several indigenous noncichlids also reappeared in the main lake, including three of the four original mormyrid species. Between 1995 and 1999, there was a dramatic increase in the proportion of haplochromines in the diet of Nile perch. When haplochromines were rare ( 1995 ), Nile perch switched from an invertebrate-dominated diet to piscivory at a large size ( 30 cm total length ). In 2000, however, Nile perch were strongly piscivorous by 5–10 cm total length. The pattern of faunal loss and recovery in Lake Nabugabo demonstrates the importance of refugia in providing the seeds of resurgence and provides a model with which to understand some changes in Lake Victoria.
Verschuren, Dirk; Johnson, Thomas C; Kling, Hedy J; Edgington, David N; Leavitt, Peter R; Brown, Erik T; Talbot, Michael R; Hecky, Robert E
Lake Victoria, the largest tropical lake in the world, suffers from severe eutrophication and the probable extinction of up to half of its 500+ species of endemic cichlid fishes. The continuing degradation of Lake Victoria's ecological functions has serious long-term consequences for the ecosystem services it provides, and may threaten social welfare in the countries bordering its shores. Evaluation of recent ecological changes in the context of aquatic food-web alterations, catchment disturbance and natural ecosystem variability has been hampered by the scarcity of historical monitoring data. Here, we present high-resolution palaeolimnological data, which show that increases in phytoplankton production developed from the 1930s onwards, which parallels human-population growth and agricultural activity in the Lake Victoria drainage basin. Dominance of bloom-forming cyanobacteria since the late 1980s coincided with a relative decline in diatom growth, which can be attributed to the seasonal depletion of dissolved silica resulting from 50 years of enhanced diatom growth and burial. Eutrophication-induced loss of deep-water oxygen started in the early 1960s, and may have contributed to the 1980s collapse of indigenous fish stocks by eliminating suitable habitat for certain deep-water cichlids. Conservation of Lake Victoria as a functioning ecosystem is contingent upon large-scale implementation of improved land-use practices. PMID:11839198
Verschuren, Dirk; Johnson, Thomas C; Kling, Hedy J; Edgington, David N; Leavitt, Peter R; Brown, Erik T; Talbot, Michael R; Hecky, Robert E
Lake Victoria, the largest tropical lake in the world, suffers from severe eutrophication and the probable extinction of up to half of its 500+ species of endemic cichlid fishes. The continuing degradation of Lake Victoria's ecological functions has serious long-term consequences for the ecosystem services it provides, and may threaten social welfare in the countries bordering its shores. Evaluation of recent ecological changes in the context of aquatic food-web alterations, catchment disturbance and natural ecosystem variability has been hampered by the scarcity of historical monitoring data. Here, we present high-resolution palaeolimnological data, which show that increases in phytoplankton production developed from the 1930s onwards, which parallels human-population growth and agricultural activity in the Lake Victoria drainage basin. Dominance of bloom-forming cyanobacteria since the late 1980s coincided with a relative decline in diatom growth, which can be attributed to the seasonal depletion of dissolved silica resulting from 50 years of enhanced diatom growth and burial. Eutrophication-induced loss of deep-water oxygen started in the early 1960s, and may have contributed to the 1980s collapse of indigenous fish stocks by eliminating suitable habitat for certain deep-water cichlids. Conservation of Lake Victoria as a functioning ecosystem is contingent upon large-scale implementation of improved land-use practices.
Carleton, Karen L; Spady, Tyrone C; Streelman, J Todd; Kidd, Michael R; McFarland, William N; Loew, Ellis R
Background Cichlid fishes have radiated into hundreds of species in the Great Lakes of Africa. Brightly colored males display on leks and vie to be chosen by females as mates. Strong discrimination by females causes differential male mating success, rapid evolution of male color patterns and, possibly, speciation. In addition to differences in color pattern, Lake Malawi cichlids also show some of the largest known shifts in visual sensitivity among closely related species. These shifts result from modulated expression of seven cone opsin genes. However, the mechanisms for this modulated expression are unknown. Results In this work, we ask whether these differences might result from changes in developmental patterning of cone opsin genes. To test this, we compared the developmental pattern of cone opsin gene expression of the Nile tilapia, Oreochromis niloticus, with that of several cichlid species from Lake Malawi. In tilapia, quantitative polymerase chain reaction showed that opsin gene expression changes dynamically from a larval gene set through a juvenile set to a final adult set. In contrast, Lake Malawi species showed one of two developmental patterns. In some species, the expressed gene set changes slowly, either retaining the larval pattern or progressing only from larval to juvenile gene sets (neoteny). In the other species, the same genes are expressed in both larvae and adults but correspond to the tilapia adult genes (direct development). Conclusion Differences in visual sensitivities among species of Lake Malawi cichlids arise through heterochronic shifts relative to the ontogenetic pattern of the tilapia outgroup. Heterochrony has previously been shown to be a powerful mechanism for change in morphological evolution. We found that altering developmental expression patterns is also an important mechanism for altering sensory systems. These resulting sensory shifts will have major impacts on visual communication and could help drive cichlid speciation
Sullivan, Melissa; Brown, Alexandria C; Clotfelter, Ethan D
Carotenoids may act as antioxidants under many circumstances. We examined the importance of carotenoids as antioxidants in the gonads of male convict cichlids (Amatitlania nigrofasciata), a species in which males lack the carotenoid-based breeding coloration that characterizes females. Male fish were fed one of four diets that included different combinations of xanthophyll and carotene carotenoids, and then we measured carotenoid concentration of the gonads, gonadosomatic index (GSI), sperm motility, and the antioxidant capacity of the gonads. Significant differences were found in gonadal carotenoid content among treatment groups, suggesting that dietary carotenoids were indeed sequestered in the gonads. There were no differences among diet groups, however, in GSI, sperm motility, or gonadal antioxidant capacity. These findings suggest that carotenoids are required only in small amounts in the testes of male convict cichlids or that they play a limited role in protecting sperm from oxidative damage.
Schwarzer, Julia; Misof, B; Schliewen, U K
Hybridization in animals is a much more common phenomenon as previously thought and may have profound implications for speciation research. The cichlid genus Steatocranus (Teleostei: Cichlidae), a close relative to members of the East African cichlid radiations, radiated under riverine conditions in the lower Congo rapids and produced a small species flock. Previous phylogenetic analyses suggested that hybridization occurred and contributed to speciation in this genus. A re-analysis of an already published 2000 loci-AFLP data set explicitly testing for patterns of ancient gene flow provided strong evidence for a highly reticulate phylogenetic history of the genus. We provide, to our knowledge, the first example of a complex reticulate network in vertebrates, including multiple closely related species connected through ancient as well as recent gene flow. In this context, the limited validity of strictly bifurcating tree hypotheses as a phylogenetic basis for hypothesis testing in evolutionary biology is discussed.
Dijkstra, Peter D.; Groothuis, Ton G. G.
It has been suggested that intrasexual competition can be a source of negative frequency-dependent selection, causing agonistic character displacement and facilitating speciation and coexistence of (sibling) species. In this paper we synthesise the evidence that male-male and female-female competition contributes to cichlid diversification, showing that competition is stronger among same-coloured individuals than those with different colours. We argue that intrasexual selection is more complex because there are several examples where males do not bias aggression towards their own type. In addition, sibling species or colour morphs often show asymmetric dominance relationships. We briefly discuss potential mechanisms that might promote the maintenance of covariance between colour and aggression-related traits even in the face of gene-flow. We close by proposing several avenues for future studies that might shed more light on the role of intrasexual competition in cichlid diversification. PMID:21785710
Background There is a critical shortage of healthcare workers in sub-Saharan Africa, and Malawi has one of the lowest physician densities in the region. One of the reasons for this shortage is inadequate retention of medical school graduates, partly due to the desire for specialization training. The University of Malawi College of Medicine has developed specialty training programs, but medical school graduates continue to report a desire to leave the country for specialization training. To understand this desire, we studied medical students’ perspectives on specialization training in Malawi. Methods We conducted semi-structured interviews of medical students in the final year of their degree program. We developed an interview guide through an iterative process, and recorded and transcribed all interviews for analysis. Two independent coders coded the manuscripts and assessed inter-coder reliability, and the authors used an “editing approach” to qualitative analysis to identify and categorize themes relating to the research aim. The University of Pittsburgh Institutional Review Board and the University of Malawi College of Medicine Research and Ethics Committee approved this study and authors obtained written informed consent from all participants. Results We interviewed 21 medical students. All students reported a desire for specialization training, with 12 (57%) students interested in specialties not currently offered in Malawi. Students discussed reasons for pursuing specialization training, impressions of specialization training in Malawi, reasons for staying or leaving Malawi to pursue specialization training and recommendations to improve training. Conclusions Graduating medical students in Malawi have mixed views of specialization training in their own country and still desire to leave Malawi to pursue further training. Training institutions in sub-Saharan Africa need to understand the needs of the country’s healthcare workforce and the needs of their
Lamboj, Anton; Trummer, Franziska; Metscher, Brian D
A new genus, Wallaceochromis, is described for the chromidotilapiine cichlids of the Pelvicachromis humilis group. It differs from Pelvicachromis sensu stricto in the number and arrangement of tubular infraorbitals, a straighter and sloping dorsal head profile with a low supraoccipital crest, higher abdominal vertebral counts (14-15 vs. 13-14), and in displaying a pattern of vertical bars on the body in several behavioral situations.
Muula, Adamson S; Maseko, Fresier C
Background The migration of health professionals from southern Africa to developed nations is negatively affecting the delivery of health care services in the source countries. Oftentimes however, it is the reasons for the out-migration that have been described in the literature. The work and domestic situations of those health professionals continuing to serve in their posts have not been adequately studied. Methods The present study utilized a qualitative data collection and analysis method. This was achieved through focus group discussions and in-depth interviews with health professionals and administrators to determine the challenges they face and the coping systems they resort to and the perceptions towards those coping methods. Results Health professionals identified the following as some of the challenges there faced: inequitable and poor remuneration, overwhelming responsibilities with limited resources, lack of a stimulating work environment, inadequate supervision, poor access to continued professionals training, limited career progression, lack of transparent recruitment and discriminatory remuneration. When asked what kept them still working in Malawi when the pressures to emigrate were there, the following were some of the ways the health professionals mentioned as useful for earning extra income to support their families: working in rural areas where life was perceived to be cheaper, working closer to home village so as to run farms, stealing drugs from health facilities, having more than one job, running small to medium scale businesses. Health professionals would also minimize expenditure by missing meals and walking to work. Conclusion Many health professionals in Malawi experience overly challenging environments. In order to survive some are involved in ethically and legally questionable activities such as receiving "gifts" from patients and pilfering drugs. The efforts by the Malawi government and the international community to retain health
District, were peculiarly killed in what was formally branded as a “ road traffic accident .” Banda had organized an in-house discussion on venturing... prevent recurrence of such rebellion. 8 THIS PAGE INTENTIONALLY LEFT BLANK 9 II. THEORETICAL FRAMEWORK Several analysts have sought to...November and AFR 36/44/92, 9 November) - Malawi: Death In Custody of Prisoner of Conscience / Fear for Physical Safety / Fear of Harassment: Orton
Henning, Frederico; Meyer, Axel
Sexual selection on male coloration is one of the main mechanisms proposed to explain the explosive speciation rates in East African cichlid fish. True eggspots are color patterns characteristic of the most species-rich lineage of cichlids, the Haplochromini, and have been suggested to be causally related to the speciation processes. Eggspots are thought to have originated by sensory exploitation and subsequently gained several roles in sexual advertisement. However, for most of these functions the evidence is equivocal. In addition, the genetic architecture of this trait still is largely unknown. We conducted bidirectional selective breeding experiments for eggspot numbers in the model cichlid, Astatotilapia burtoni. After two generations, low lines responded significantly, whereas the high lines did not. Body size was both phenotypically and genotypically correlated with eggspot number and showed correlated response to selection. Males with higher numbers of eggspots were found to sire larger offspring. Despite the potential to act as honest indicators of fitness, the behavioral experiments showed no evidence of a role in either intra- or inter-sexual selection. Visual-based female preference was instead explained by courtship intensity. The evolution of this trait has been interpreted in light of adaptive theories of sexual selection, however the present and published results suggest the influence of non-adaptive factors such as sensory exploitation, environmental constraints and sexual antagonism. PMID:22937082
Willis, Stuart; Watkins, Crystal; Honeycutt, Rodney L.; Winemiller, Kirk O.
Understanding of relationships between morphology and ecological performance can help to reveal how natural selection drives biological diversification. We investigate relationships between feeding behavior, foraging performance and morphology within a diverse group of teleost fishes, and examine the extent to which associations can be explained by evolutionary relatedness. Morphological adaptation associated with sediment sifting was examined using a phylogenetic linear discriminant analysis on a set of ecomorphological traits from 27 species of Neotropical cichlids. For most sifting taxa, feeding behavior could be effectively predicted by a linear discriminant function of ecomorphology across multiple clades of sediment sifters, and this pattern could not be explained by shared evolutionary history alone. Additionally, we tested foraging efficiency in seven Neotropical cichlid species, five of which are specialized benthic feeders with differing head morphology. Efficiency was evaluated based on the degree to which invertebrate prey could be retrieved at different depths of sediment. Feeding performance was compared both with respect to feeding mode and species using a phylogenetic ANCOVA, with substrate depth as a covariate. Benthic foraging performance was constant across sediment depths in non-sifters but declined with depth in sifters. The non-sifting Hypsophrys used sweeping motions of the body and fins to excavate large pits to uncover prey; this tactic was more efficient for consuming deeply buried invertebrates than observed among sediment sifters. Findings indicate that similar feeding performance among sediment-sifting cichlids extracting invertebrate prey from shallow sediment layers reflects constraints associated with functional morphology and, to a lesser extent, phylogeny. PMID:24603485
Fabrin, T M C; Gasques, L S; Prioli, S M A P; Prioli, A J
The use of molecular markers has contributed to phylogeny and to the reconstruction of species' evolutionary history. Each region of the genome has different evolution rates, which may or may not identify phylogenetic signal at different levels. Therefore, it is important to assess new molecular markers that can be used for phylogenetic reconstruction. Regions that may be associated with species characteristics and are subject to selective pressure, such as opsin genes, which encode proteins related to the visual system and are widely expressed by Cichlidae family members, are interesting. Our aim was to identify a new nuclear molecular marker that could establish the phylogeny of Neotropical cichlids and is potentially correlated with the visual system. We used Bayesian inference and maximum likelihood analysis to support the use of the nuclear opsin LWS gene in the phylogeny of eight Neotropical cichlid species. Their use concatenated to the mitochondrial gene COI was also tested. The LWS gene fragment comprised the exon 2-4 region, including the introns. The LWS gene provided good support for both analyses up to the genus level, distinguishing the studied species, and when concatenated to the COI gene, there was a good support up to the species level. Another benefit of utilizing this region, is that some polymorphisms are associated with changes in spectral properties of the LWS opsin protein, which constitutes the visual pigment that absorbs red light. Thus, utilization of this gene as a molecular marker to study the phylogeny of Neotropical cichlids is promising.
Lehtonen, Topi K; Sowersby, Will; Gagnon, Karine; Wong, Bob B M
The ability to assess the threat posed by competitors, and to respond appropriately, is important for reducing the costs of aggression. In this respect, aggression directed toward heterospecifics is often just as significant as aggression among conspecifics. This is especially true for cichlid fish that share breeding grounds with heterospecifics. Indeed, cichlids are known to differentiate not only between conspecifics that pose different levels of threat but also between heterospecific territorial intruders by directing more aggression toward nonbreeding individuals. To assess whether the ability to make such distinctions could be based on color cues alone, we carried out a field study in which we experimentally presented Amphilophus sagittae cichlid pairs with model intruders of a sympatric congener, Amphilophus xiloaensis, in breeding versus nonbreeding coloration. Consistent with our prediction, we found that A. sagittae exhibited more aggression toward A. xiloaensis models of the latter color type. The results are, to our knowledge, the first to show that territory holders can, based on coloration alone, assess variation among individuals of a species other than their own in the threat posed to offspring survival.
Mohammadi, F; Mousavi, S. M.; Ahmadmoradi, E.; Zakeri, M.; Jahedi, A.
Using probiotics can control pathogens by a variety of mechanisms. Probiotics can promote growth performance and have, therefore, become increasingly important in the aquaculture industry. Convict Cichlid belongs to the family of Cichlidae and is known for its rapid development in laboratory conditions and is suitable for behavioral examinations. The aim of this study was to evaluate the effects of Saccharomyces cerevisiae on growth performance, survival rate and body composition of Convict Cichlids (Amatitlania nigrofasciata). One hundred sixty eight Convict Cichlids (mean weight: 2.1 ± 0.12 g and mean length: 2.2 ± 0.05 cm) were fed by commercial diets with different concentrations of S. cerevisiae (0, 0.5%, 1%, 2%). At the end of the experiment, survival rate and growth indices were measured. Based on the results, growth performance significantly increased with probiotic, S. cerevisiae, specially, at the 2% probiotic level of concentration. In the present study, the best FCR (feed conversion rate), SGR (specific growth rate), CF (condition factor) and BWG (body weight gain) values were observed in a 2% concentration of S. cerevisiae. The results suggest that this yeast could improve feed utilization in this fish species. PMID:27175152
Mohammadi, F; Mousavi, S M; Ahmadmoradi, E; Zakeri, M; Jahedi, A
Using probiotics can control pathogens by a variety of mechanisms. Probiotics can promote growth performance and have, therefore, become increasingly important in the aquaculture industry. Convict Cichlid belongs to the family of Cichlidae and is known for its rapid development in laboratory conditions and is suitable for behavioral examinations. The aim of this study was to evaluate the effects of Saccharomyces cerevisiae on growth performance, survival rate and body composition of Convict Cichlids (Amatitlania nigrofasciata). One hundred sixty eight Convict Cichlids (mean weight: 2.1 ± 0.12 g and mean length: 2.2 ± 0.05 cm) were fed by commercial diets with different concentrations of S. cerevisiae (0, 0.5%, 1%, 2%). At the end of the experiment, survival rate and growth indices were measured. Based on the results, growth performance significantly increased with probiotic, S. cerevisiae, specially, at the 2% probiotic level of concentration. In the present study, the best FCR (feed conversion rate), SGR (specific growth rate), CF (condition factor) and BWG (body weight gain) values were observed in a 2% concentration of S. cerevisiae. The results suggest that this yeast could improve feed utilization in this fish species.
Schneider, Ralf F; Li, Yuanhao; Meyer, Axel; Gunter, Helen M
Phenotypic plasticity is the ability of organisms with a given genotype to develop different phenotypes according to environmental stimuli, resulting in individuals that are better adapted to local conditions. In spite of their ecological importance, the developmental regulatory networks underlying plastic phenotypes often remain uncharacterized. We examined the regulatory basis of diet-induced plasticity in the lower pharyngeal jaw (LPJ) of the cichlid fish Astatoreochromis alluaudi, a model species in the study of adaptive plasticity. Through raising juvenile A. alluaudi on either a hard or soft diet (hard-shelled or pulverized snails) for between 1 and 8 months, we gained insight into the temporal regulation of 19 previously identified candidate genes during the early stages of plasticity development. Plasticity in LPJ morphology was first detected between 3 and 5 months of diet treatment. The candidate genes, belonging to various functional categories, displayed dynamic expression patterns that consistently preceded the onset of morphological divergence and putatively contribute to the initiation of the plastic phenotypes. Within functional categories, we observed striking co-expression, and transcription factor binding site analysis was used to examine the prospective basis of their coregulation. We propose a regulatory network of LPJ plasticity in cichlids, presenting evidence for regulatory crosstalk between bone and muscle tissues, which putatively facilitates the development of this highly integrated trait. Through incorporating a developmental time-course into a phenotypic plasticity study, we have identified an interconnected, environmentally responsive regulatory network that shapes the development of plasticity in a key innovation of East African cichlids.
Langen, Kathrin; Thünken, Timo; Bakker, Theo C. M.
Alternative reproductive tactics are widespread in fishes, increasing the potential for sperm competition. Sperm competition has enormous impact on both variation in sperm numbers and sperm size. In cichlids, the sperm competition risk is very divergent and longer sperm are usually interpreted as adaptation to sperm competition. Here we examined whether sneaking tactics exist in Pelvicachromis taeniatus, a socially monogamous cichlid with biparental brood care from West Africa. The small testis indicates low gonadal investment which is typical for genetically monogamous species. In contrast, sperm length with up to 85 μm is extraordinarily long. We examined the reproductive behaviour of ten groups with a male-biased sex ratio under semi-natural conditions via continuous video recording. We recorded spawning site preferences and correlates of reproductive success and conducted paternity tests using microsatellites. Safe breeding sites that could be successfully defended were preferred. All offspring could be assigned to their parents and no multiple paternities were detected. Body size of spawning pairs predicted their spawning probability and offspring hatching rate suggesting benefits from mating with large individuals. Our study suggests low risk of sperm competition under the given conditions in P. taeniatus and thus first evidence for genetic monogamy in a substrate breeding cichlid. PMID:24386589
O'Hare, B; Curtis, M
This analysis examines the gaps in health care financing in Malawi and how foregone taxes could fill these gaps. It begins with an assessment of the disease burden and government health expenditure. Then it analyses the tax revenues foregone by the government of Malawi by two main routes: Illicit financial flows (IFF) from the country, Tax incentives. We find that there are significant financing gaps in the health sector; for example, government expenditure is United States Dollars (USD) 177 million for 2013/2014 while projected donor contribution in 2013/2014 is USD 207 million and the total cost for the minimal health package is USD 535 million. Thus the funding gap between the government budget for health and the required spending to provide the minimal package for 2013/2014 is USD 358 million. On the other hand we estimate that almost USD 400 million is lost through IFF and corporate utilization of tax incentives each year. The revenues foregone plus the current government health spending would be sufficient to cover the minimal public health package for all Malawians and would help tackle Malawi's disease burden. Every effort must be made, including improving transparency and revising laws, to curtail IFF and moderate tax incentives.
Water insecurity is one of the most pressing issues currently faced by Malawi. The consequences of these issues are borne significantly by women, who are most directly involved with water provision and use, particularly at the household level. Since the mid-1990s, Malawi has undertaken a process of water policy reform. Reflective of international…
Kadzamira, Esme Chipo; Kunje, Demis
A study was commissioned by the United States Agency for International Development (USAID), Bureau for Africa, to investigate the changing roles of nongovernmental organizations (NGOs) in basic education in Africa. This Malawi case study was conducted by the Centre for Educational Research and Training of the University of Malawi. Information for…
Kalanda, Boniface Francis
Malawi is one of the countries with a youthful population. Youths are susceptible to various social-economic pressures that put their well being into jeopardy. One of the issues that affect youth is early drop out of school, drug abuse and contracting sexually transmitted diseases including HIV. To address these issues, the Malawi Government…
Lupafya, Phindile Chitsulo; Mwagomba, Beatrice L. Matanje; Hosig, Kathy; Maseko, Lucy M.; Chimbali, Henry
Malawi is a Sub-Saharan African country experiencing the epidemiological transition from predominantly infectious to noncommunicable diseases (NCDs) with dramatically increasing prevalence of lifestyle-related diseases such as obesity, hypertension, and diabetes. Malawi's 2011-2016 Health Sector Strategic Plan included NCDs, and an NCD Control…
Paperno, R.; Ruiz-Carus, R.; Krebs, J.M.; McIvor, C.C.
Introduced exotic species are a well-recognized problem in Florida's subtropical ecosystems. The presence of the exotic Mayan cichlid (Cichlasoma urophthalmus) was first confirmed in Florida in 1983, when numerous individuals were found in the northeastern Florida Bay. Since then, this species has continued to expand its range northward. The capture, beginning in October 2004 to present, of large numbers of Mayan cichlids from central Florida's east- and west-coast mangrove systems north of 28°N latitude is documented here. Mayan cichlids in a wide range of sizes (estimated ages 0-7 years) at both east- and west-coast sites were collected. In addition, macroscopic examination of gonads showed the presence of developing eggs. The occurrence of multiple age-classes, maturing individuals, cichlid nests, and juveniles, plus repeated collections over a four-year period, indicates that the Mayan cichlid is successfully reproducing and surviving the average winter temperatures in some estuarine waters in central Florida.
Machado-Schiaffino, Gonzalo; Henning, Frederico; Meyer, Axel
The spectacular species richness of cichlids and their diversity in morphology, coloration, and behavior have made them an ideal model for the study of speciation and adaptive evolution. Hypertrophic lips evolved repeatedly and independently in African and Neotropical cichlid radiations. Cichlids with hypertrophic lips forage predominantly in rocky crevices and it has been hypothesized that mechanical stress caused by friction could result in larger lips through phenotypic plasticity. To test the influence of the environment on the size and development of lips, we conducted a series of breeding and feeding experiments on Midas cichlids. Full-sibs of Amphilophus labiatus (thick-lipped) and Amphilophus citrinellus (thin-lipped) each were split into a control group which was fed food from the water column and a treatment group whose food was fixed to substrates. We found strong evidence for phenotypic plasticity on lip area in the thick-lipped species, but not in the thin-lipped species. Intermediate phenotypic values were observed in hybrids from thick- and thin-lipped species reared under "control" conditions. Thus, both a genetic, but also a phenotypic plastic component is involved in the development of hypertrophic lips in Neotropical cichlids. Moreover, species-specific adaptive phenotypic plasticity was found, suggesting that plasticity is selected for in recent thick-lipped species.
Kim, Sung Wook; Skordis-Worrall, Jolene; Haghparast-Bidgoli, Hassan; Pulkki-Brännström, Anni-Maria
Background Human immunodeficiency virus (HIV) is a significant contributor to Malawi's burden of disease. Despite a number of studies describing socio-economic differences in HIV prevalence, there is a paucity of evidence on socio-economic inequity in HIV testing in Malawi. Objective To assess horizontal inequity (HI) in HIV testing in Malawi. Design Data from the Demographic and Health Surveys (DHSs) 2004 and 2010 in Malawi are used for the analysis. The sample size for DHS 2004 was 14,571 (women =11,362 and men=3,209), and for DHS 2010 it was 29,830 (women=22,716 and men=7,114). The concentration index is used to quantify the amount of socio-economic-related inequality in HIV testing. The inequality is a primary method in this study. Corrected need, a further adjustment of the standard decomposition index, was calculated. Standard HI was compared with corrected need-adjusted inequity. Variables used to measure health need include symptoms of sexually transmitted infections. Non-need variables include wealth, education, literacy and marital status. Results Between 2004 and 2010, the proportion of the population ever tested for HIV increased from 15 to 75% among women and from 16 to 54% among men. The need for HIV testing among men was concentrated among the relatively wealthy in 2004, but the need was more equitably distributed in 2010. Standard HI was 0.152 in 2004 and 0.008 in 2010 among women, and 0.186 in 2004 and 0.04 in 2010 among men. Rural–urban inequity also fell in this period, but HIV testing remained pro-rich among rural men (HI 0.041). The main social contributors to inequity in HIV testing were wealth in 2004 and education in 2010. Conclusions Inequity in HIV testing in Malawi decreased between 2004 and 2010. This may be due to the increased support to HIV testing by global donors over this period. PMID:27790970
Anderson, Caleb; Werdenig, Alexandra; Koblmüller, Stephan; Sefc, Kristina M
Mating system variability is known to exist between and within species, often due to environmental influences. An open question is whether, vice versa, similar environmental conditions entail congruent mating behavior, for example in terms of multiple paternity, in species or populations sharing largely comparable breeding modes. This study employed microsatellite markers to investigate the incidence of multiple paternity in Cyprichromis coloratus and Cyprichromis leptosoma, two sympatric, closely related, mouthbrooding Lake Tanganyika cichlids with similar ecological and behavioral characteristics including the formation of open-water schools. Mouthbrooding females of both species were collected from the same mixed-species breeding school at the same time, minimizing environmental variation during courtship and mating. In C. coloratus, four of 12 broods had more than one sire, with a mean of 1.33 reconstructed sires per brood. C. leptosoma exhibited multiple paternity in 18 of 22 broods, with a mean of 2.59 or 2.86 reconstructed sires per brood according to the programs gerud and colony, respectively. In addition, two broods were found to contain offspring transplanted from another brood. There was no significant difference in brood size between species, but mean sire number did differ significantly. Hence, substantial similarity in reproductive behavior along with shared environmental conditions during courtship and spawning did not lead to equal rates of polyandry or sneaking in the two species.
Hablützel, P I; Vanhove, M P M; Grégoir, A F; Hellemans, B; Volckaert, F A M; Raeymaekers, J A M
Studying the genetic basis of host-parasite interactions represents an outstanding opportunity to observe eco-evolutionary processes. Established candidates for such studies in vertebrates are immunogenes of the major histocompatibility complex (MHC). The MHC has been reported to reach high intra- and interindividual diversity, and a diverse MHC might be advantageous when facing infections from multiple parasites. However, other studies indicated that individuals with an intermediate number of MHC alleles are less infected with parasites or have other fitness advantages. In this study, we assessed the optimal number of MHC alleles in the blunt-head cichlid Tropheus moorii from Lake Tanganyika. We investigated the influence of the interindividual variation in number of MHC length variants on parasite infection and body condition, measured by the amount of perivisceral fat reserves. Surprisingly, there was no correlation between parasite infection and number of MHC length variants or perivisceral fat deposits. However, the individual number of MHC length variants significantly correlated with the amount of perivisceral fat deposits in males, suggesting that male individuals with an intermediate number of alleles might be able to use their fat reserves more efficiently.
Redescription of Neoechinorhynchus (Neoechinorhynchus) golvani Salgado-Maldonado, 1978 (Acanthocephala: Neoechinorhynchidae) and description of a new species from freshwater cichlids (Teleostei: Cichlidae) in Mexico.
A redescription of Neoechinorhynchus (Neoechinorhynchus) golvani Salgado-Maldonado (An Inst Biol Univ Nal Autón Méx, Ser Zool 49:35-47, 1978) is presented, based on adult specimens collected from the type host Paraneetroplus fenestratus from the type location, the Lago de Catemaco lake, Veracruz state, Mexico, and its presence is recorded in other cichlids. Detailed studies of N. (N.) golvani using light microscopy revealed some taxonomically important, previously unreported features, such as the size and shape of fully developed adult males and females, and the structure of the eggs. Morphological variability in N. (N.) golvani is described. Based on these data, the geographic distribution of this species is documented. Neoechinorhynchus (Neoechinorhynchus) panucensis n. sp. is described from Herichthys labridens (Pellegrin), Amatitlania nigrofasciata (Günther), and Herichthys cyanoguttatus Baird and Girard (all of them Cichlidae), collected in the Río Atlapexco, a tributary to the upper Río Panuco basin, Hidalgo State, Mexico. This new species stand up alone because of its minute proboscis (♂ 50 × 60, ♀ 42-55 (48.5) × 48-63 (57.7)) and anterior hooks (♂ 27-30 (28.8) × 3-5 (4), ♀ 28-32 (30) × 5 (5)). A key to the species of Neoechinorhynchus recorded from freshwater fishes in Central and South America is included.
Ohrn, Deborah Gore, Ed.
This quarterly publication of the State Historical Society of Iowa features articles and activities for elementary school students. This summer issue focuses on the topic of lake life. The issue includes the following features: (1) "Where the Lakes Are Map"; (2) "Letter from the Lake"; (3) "Lake People"; (4)…
Singogo, E; Kanike, E; van Lettow, M; Cataldo, F; Zachariah, R; Bissell, K; Harries, A D
Paper-based village registers were introduced 5 years ago in Malawi as a tool to measure vital statistics of births and deaths at the population level. However, usage, completeness and accuracy of their content have never been formally evaluated. In Traditional Authority Mwambo, Zomba district, Malawi, we assessed 280 of the 325 village registers with respect to (i) characteristics of village headmen who used village registers, (ii) use and content of village registers, and (iii) whether village registers provided accurate information on births and deaths. All village headpersons used registers. There were 185 (66%) registers that were regarded as 95% completed, and according to the registers, there were 115 840 people living in the villages in the catchment area. In 2011, there were 1753 births recorded in village registers, while 6397 births were recorded in health centre registers in the same catchment area. For the same year, 199 deaths were recorded in village registers, giving crude death rates per 100 000 population of 189 for males and 153 for females. These could not be compared with death rates in health centre registers due to poor and inconsistent recording in these registers, but they were compared with death rates obtained from the 2010 Malawi Demographic Health Survey that reported 880 and 840 per 100 000 for males and females, respectively. In conclusion, this study shows that village registers are a potential source for vital statistics. However, considerable inputs are needed to improve accuracy of births and deaths, and there are no functional systems for the collation and analysis of data at the traditional authority level. Innovative ways to address these challenges are discussed, including the use of solar-powered electronic village registers and mobile phones, connected with each other and the health facilities and the District Commissioner's office through the cellular network and wireless coverage.
Pierotti, Rachael S.; Young-DeMarco, Linda; Watkins, Susan
This paper examines the extent to which developmental idealism has been disseminated in Malawi. Developmental idealism is a set of beliefs and values about development and the relationships between development and family structures and behavior. Developmental idealism states that attributes of societies and families defined as modern are better than attributes defined as traditional, that modern societies help produce modern families, that modern families facilitate the achievement of modern societies, and that the future will bring family change in the direction of modernity. Previous research has demonstrated that knowledge of developmental idealism is widespread in many places around the world, but provides little systematic data about it in sub-Saharan Africa or how knowledge of it is associated with certain demographic characteristics in that region. In this paper, we address this issue by examining whether ordinary people in two settings in Malawi, a sub-Saharan African country, have received and understood messages that are intended to associate development with certain types of family forms and family behaviors. We then examine associations between demographic characteristics and developmental idealism to investigate possible mechanisms linking global discourse about development to the grassroots. We analyze data collected in face-to-face surveys from two samples of Malawian men in 2009 and 2010, one rural, the other in a low-to-medium income neighborhood of a city. Our analysis of these survey data shows considerable evidence that many developmental idealism beliefs have been spread in that country and that education has positive effects on beliefs in the association between development and family attributes. We also find higher levels of developmental idealism awareness in the urban sample than we do in the rural sample, but once dissimilarities in education and wealth between the two samples are controlled, awareness levels no longer differed between
Mueller, Dirk H.; Lungu, Douglas; Acharya, Arnab; Palmer, Natasha
Increasingly seen as a useful tool of health policy, Essential or Minimal Health Packages direct resources to interventions that aim to address the local burden of disease and be cost-effective. Less attention has been paid to the delivery mechanisms for such interventions. This study aimed to assess the degree to which the Essential Health Package (EHP) in Malawi was available to its population and what health system constraints impeded its full implementation. The first phase of this study comprised a survey of all facilities in three districts including interviews with all managers and clinical staff. In the second and third phase, results were discussed with District Health Management Teams and national level stakeholders, respectively, including representatives of the Ministry of Health, Central Medical Stores, donors and NGOs. The EHP in Malawi is focussing on the local burden of disease; however, key constraints to its successful implementation included a widespread shortage of staff due to vacancies but also caused by frequent trainings and meetings (only 48% of expected man days of clinical staff were available; training and meetings represented 57% of all absences in health centres). Despite the training, the percentage of health workers aware of vital diagnostic and therapeutic approaches to EHP conditions was weak. Another major constraint was shortages of vital drugs at all levels of facilities (e.g. Cotrimoxazole was sufficiently available to treat the average number of patients in only 27% of health centres). Although a few health workers noted some improvement in infrastructure and working conditions, they still considered them to be widely inadequate. In Malawi, as in similar resource poor countries, greater attention needs to be given to the health system constraints to delivering health care. Removal of these constraints should receive priority over the considerable focus on the development and implementation of essential packages of
Maseko, Fresier C.; Chirwa, Maureen L.; Muula, Adamson S.
Background Cervical cancer remains the leading cause of cancer death among women in sub-Saharan Africa. In Malawi, very few women have undergone screening and the incidence of cervical cancer is on the increase as is the case in most developing countries. We aimed at exploring and documenting health system gaps responsible for the poor performance of the cervical cancer prevention program in Malawi. Design The study was carried out in 14 randomly selected districts of the 29 districts of Malawi. All cervical cancer service providers in these districts were invited to participate. Two semi-structured questionnaires were used, one for the district cervical cancer coordinators and the other for the service providers. The themes of both questionnaires were based on World Health Organization (WHO) health system frameworks. A checklist was also developed to audit medical supplies and equipment in the cervical cancer screening facilities. The two questionnaires together with the medical supplies and equipment checklist were piloted in Chikwawa district before being used as data collection tools in the study. Quantitative data were analyzed using STATA and qualitative in NVIVO. Results Forty-one service providers from 21 health facilities and 9 district coordinators participated in the study. Our findings show numerous health system challenges mainly in areas of health workforce and essential medical products and technologies. Seven out of the 21 health facilities provided both screening and treatment. Results showed challenges in the management of the cervical cancer program at district level; inadequate service providers who are poorly supervised; lack of basic equipment and stock-outs of basic medical supplies in some health facilities; and inadequate funding of the program. In most of the health facilities, services providers were not aware of the policy which govern their work and that they did not have standards and guidelines for cervical cancer screening and
The Malawi Social Action Fund (MASAF) is implementing a 12 year programme to close service gaps in rural communities. These service gaps are primarily those in health, education, household food security, water and sanitation, transport and communications. The impact indicators of the Project are selected Millennium Development Goal indicators. MASAF conducted a baseline study of the MDG indicators for all districts in Malawi. This paper presents available health related MDG baseline indicators for all districts in Malawi. Other stakeholders implementing health interventions could use these baseline indicators for planning purposes.
O'Connor, Constance M.; Reddon, Adam R.; Marsh-Rollo, Susan E.; Hellmann, Jennifer K.; Ligocki, Isaac Y.; Hamilton, Ian M.; Balshine, Sigal
Social interactions facilitate pathogen transmission and increase virulence. Therefore, species that live in social groups are predicted to suffer a higher pathogen burden, to invest more heavily in immune defence against pathogens, or both. However, there are few empirical tests of whether social species indeed invest more heavily in immune defence than non-social species. In the current study, we conducted a phylogenetically controlled comparison of innate immune response in Lamprologine cichlid fishes. We focused on three species of highly social cichlids that live in permanent groups and exhibit cooperative breeding ( Julidochromis ornatus, Neolamprologus pulcher and Neolamprologus savoryi) and three species of non-social cichlids that exhibit neither grouping nor cooperative behaviour ( Telmatochromis temporalis, Neolamprologus tetracanthus and Neolamprologus modestus). We quantified the innate immune response by injecting wild fishes with phytohaemagglutinin (PHA), a lectin that causes a cell-mediated immune response. We predicted that the three highly social species would show a greater immune reaction to the PHA treatment, indicating higher investment in immune defence against parasites relative to the three non-social species. We found significant species-level variation in immune response, but contrary to our prediction, this variation did not correspond to social system. However, we found that immune response was correlated with territory size across the six species. Our results indicate that the common assumption of a positive relationship between social system and investment in immune function may be overly simplistic. We suggest that factors such as rates of both in-group and out-group social interactions are likely to be important mediators of the relationship between sociality and immune function.
Fishes have evolved an astonishing diversity of peripheral (accessory/ancillary) auditory structures to improve hearing based on their ability to transmit oscillations of gas bladder walls to the inner ears. So far it is unclear to what degree the size of the bladder and the linkage to the ear affect hearing in fishes. An interfamilial study in catfishes revealed that families which possess large, single swim bladders and one to four Weberian ossicles were more sensitive at higher frequencies (≥1 kHz) than families which have small, paired, and encapsulated bladders and one to two ossicles. An intrafamilial investigation in thorny catfishes (family Doradidae) revealed that small differences in bladder morphology did not affect hearing similarly. Members of the cichlid family possess an even larger variation in peripheral auditory structures than catfishes. The linkage between the swim bladder and ear can either be present via anterior extensions of the bladder or be completely absent (in contrast to catfishes). Representatives having large bladders with extensions had the best sensitivities. Cichlids lacking extensions had lower sensitivities above 0.3 kHz. Species with a vestigial swim bladder exhibited a smaller hearing bandwidth than those with larger swim bladder (maximum frequency: 0.7 kHz vs. 3 kHz). Catfishes and cichlids reveal that larger gas bladders and more pronounced connections between the swim bladder and the inner ear result in improved hearing at higher frequencies. The lack of a connection between a large bladder and the inner ear does not necessarily result in a smaller detectable frequency range.
Albertson, R Craig; Powder, Kara E; Hu, Yinan; Coyle, Kaitlin P; Roberts, Reade B; Parsons, Kevin J
Variation in pigmentation type and levels is a hallmark of myriad evolutionary radiations, and biologists have long been fascinated by the factors that promote and maintain variation in coloration across populations. Here, we provide insights into the genetic basis of complex and continuous patterns of colour variation in cichlid fishes, which offer a vast diversity of pigmentation patterns that have evolved in response to both natural and sexual selection. Specifically, we crossed two divergent cichlid species to generate an F2 mapping population that exhibited extensive variation in pigmentation levels and patterns. Our experimental design is robust in that it combines traditional quantitative trait locus (QTL) analysis with population genomics, which has allowed us to move efficiently from QTL interval to candidate gene. In total, we detected 41 QTL and 13 epistatic interactions that underlie melanocyte- and xanthophore-based coloration across the fins and flanks of these fishes. We also identified 2 QTL and 1 interaction for variation in the magnitude of integration among these colour traits. This finding in particular is notable as there are marked differences both within and between species with respect to the complexity of pigmentation patterns. While certain individuals are characterized by more uniform 'integrated' colour patterns, others exhibit many more degrees of freedom with respect to the distribution of colour 'modules' across the fins and flank. Our data reveal, for the first time, a genetic basis for this difference. Finally, we implicate pax3a as a mediator of continuous variation in the levels of xanthophore-based colour along the cichlid flank.
Beier, M.; Hilbig, R.; Anken, R.
Inner ear otolith growth in terms of mineralisation mainly depends on the enzyme carbonic anhydrase (CAH). CAH is located in specialised, mitochondria-rich macular cells (ionocytes), which are involved in the endolymphatic ion exchange, and the enzyme is responsible for the provision of the pH-value necessary for otolithic calcium carbonate deposition. In the present study, for the first time the localisation of histochemically demonstrated CAH was analysed during the early larval development of a teleost, the cichlid fish Oreochromis mossambicus. CAH-reactivity was observed already in stage 7 animals (onset of otocyst development; staging follows Anken et al. [Anken, R., Kappel, T., Slenzka, K., Rahmann, H. The early morphogenetic development of the cichlid fish, Oreochromis mossambicus (Perciformes, Teleostei). Zool. Anz. 231, 1-10, 1993]). Neuroblasts (from which sensory and supporting cells are derived) proved to be CAH-positive. Already at stage 12 (hatch), CAH-positive regions could be attributed to ionocyte containing regions both in the so-called meshwork and patches area of the macula (i.e., clearly before ionocytes can be identified on ultrastructural level or by employing immunocytochemistry). In contrast to the circumstances observed in mammalian species, sensory hair cells stained negative for CAH in the cichlid. With the onset of stage 16 (finray primordia in dorsal fin, yolk-sac being increasingly absorbed), CAH-reactivity was observed in the vestibular nerve. This indicates the onset of myelinisation and thus commencement of operation. The localisation of CAH in the inner ear of fish (especially the differences in comparison to mammals) is discussed on the basis of its role in otolith calcification. Since the vestibular system is a detector of acceleration and thus gravity, also aspects regarding effects of altered gravity on CAH and hence on the mineralisation of otoliths in an adaptive process are addressed.
McLaughlin, Kirsty Elizabeth; Kunc, Hansjoerg P
Anthropogenic noise can affect behaviour across a wide range of species in both terrestrial and aquatic environments. However, behaviours might not be affected in isolation. Therefore, a more holistic approach investigating how environmental stressors, such as noise pollution, affect different behaviours in concert is necessary. Using tank-based noise exposure experiments, we tested how changes in the acoustic environment affect the behaviour of the cichlid Amatitlania nigrofasciata. We found that exposure to anthropogenic noise affected a couple of behaviours: an increase in sheltering was accompanied by a decrease in foraging. Our results highlight the multiple negative effects of an environmental stressor on an individual's behaviour.
Swerdlow, D. L.; Malenga, G.; Begkoyian, G.; Nyangulu, D.; Toole, M.; Waldman, R. J.; Puhr, D. N.; Tauxe, R. V.
Between 23 August and 15 December 1990 an epidemic of cholera affected Mozambican refugees in Malawi causing 1931 cases (attack rate = 2.4%); 86% of patients had arrived in Malawi < 3 months before illness onset. There were 68 deaths (case-fatality rate = 3.5%); most deaths (63%) occurred within 24 h of hospital admission which may have indicated delayed presentation to health facilities and inadequate early rehydration. Mortality was higher in children < 4 years old and febrile deaths may have been associated with prolonged i.v. use. Significant risk factors for illness (P < 0.05) in two case-control studies included drinking river water (odds ratio [OR] = 3.0); placing hands into stored household drinking water (OR = 6.0); and among those without adequate firewood to reheat food, eating leftover cooked peas (OR = 8.0). Toxigenic V. cholerae O1, serotype Inaba, was isolated from patients and stored household water. The rapidity with which newly arrived refugees became infected precluded effective use of a cholera vaccine to prevent cases unless vaccination had occurred immediately upon camp arrival. Improved access to treatment and care of paediatric patients, and increased use of oral rehydration therapy, could decrease mortality. Preventing future cholera outbreaks in Africa will depend on interrupting both waterborne and foodborne transmission of this pathogen. PMID:9207730
Chimombo, Joseph Patrick Godson
Free primary education was introduced to Malawi in 1994, and it led to an increase in enrolment of one million pupils. The present contribution gathers case studies of 10 schools which sought information on the general conditions of communities and schools as well as on enrolment, resources, staffing, absenteeism etc. The results highlight the difficulties of implementing Education for All (EFA). Generally speaking, schools have been poorly supplied by the state. Effectively, many pupils have been marshaled into schools not properly equipped in terms of material and human resources. As a result, EFA in Malawi has been undermined by a persistently large number of pupils who drop out before attaining permanent literacy. Yet providing quality education is more than just a question of resources. From the points of view of parents and pupils, quality education requires relevance to local needs; adaptability to local conditions; and flexibility in addressing cultural obstacles. Overall, as is shown here, it appears that increased access to schooling has been achieved at the expense of the quality of education offered.
Kaphagawani, Nanzen Caroline; Kalipeni, Ezekiel
This study explores sociocultural and other risk factors associated with unplanned teenage pregnancy in Zomba district of Malawi. Data were obtained from 505 participants under the age of 20 years using a questionnaire administered through face-to-face interviews held at five antenatal clinics. The data were analysed using descriptive statistics, frequency tables and chi-square analysis which allowed comparative understanding of the sociocultural risk factors for planned and unplanned teenage pregnancy in Zomba district. The findings revealed that teenage pregnancy is a major health and social problem. Over 76% of the teenage respondents in the study had experienced unplanned pregnancy. Among the prominent factors that stood out in the analysis for this high rate of teenage pregnancy were early sex and marriage, low contraceptive use, low educational levels, low socio-economic status, lack of knowledge of reproductive and sexual health, gender inequity, and physical/sexual violence. The consequences on teenage mothers of unplanned pregnancy have been tragic and have compromised their physical, psychological and socioeconomic wellbeing, not just on them but also their families and society at large. The findings point to the need for a multi-sectoral approach to tackle the problem on teenage pregnancy in this district, and likely throughout Malawi.
Dougill, Andrew J; Whitfield, Stephen; Stringer, Lindsay C; Vincent, Katharine; Wood, Benjamin T; Chinseu, Edna L; Steward, Peter; Mkwambisi, David D
Conservation agriculture (CA) practices of reduced soil tillage, permanent organic soil coverage and intercropping/crop rotation, are being advocated globally, based on perceived benefits for crop yields, soil carbon storage, weed suppression, reduced soil erosion and improved soil water retention. However, some have questioned their efficacy due to uncertainty around the performance and trade-offs associated with CA practices, and their compatibility with the diverse livelihood strategies and varied agro-ecological conditions across African smallholder systems. This paper assesses the role of key institutions in Malawi in shaping pathways towards more sustainable land management based on CA by outlining their impact on national policy-making and the design and implementation of agricultural development projects. It draws on interviews at national, district and project levels and a multi-stakeholder workshop that mapped the institutional landscape of decision-making for agricultural land management practices. Findings identify knowledge gaps and institutional barriers that influence land management decision-making and constrain CA uptake. We use our findings to set out an integrated roadmap of research needs and policy options aimed at supporting CA as a route to enhanced sustainable land management in Malawi. Findings offer lessons that can inform design, planning and implementation of CA projects, and identify the multi-level institutional support structures required for mainstreaming sustainable land management in sub-Saharan Africa.
Hurst, Rachel; Siyame, Edwin W P; Young, Scott D; Chilimba, Allan D C; Joy, Edward J M; Black, Colin R; Ander, E Louise; Watts, Michael J; Chilima, Benson; Gondwe, Jellita; Kang'ombe, Dalitso; Stein, Alexander J; Fairweather-Tait, Susan J; Gibson, Rosalind S; Kalimbira, Alexander A; Broadley, Martin R
Selenium (Se) is an essential human micronutrient with critical roles in immune functioning and antioxidant defence. Estimates of dietary Se intakes and status are scarce for Africa although crop surveys indicate deficiency is probably widespread in Malawi. Here we show that Se deficiency is likely endemic in Malawi based on the Se status of adults consuming food from contrasting soil types. These data are consistent with food balance sheets and composition tables revealing that >80% of the Malawi population is at risk of dietary Se inadequacy. Risk of dietary Se inadequacy is >60% in seven other countries in Southern Africa, and 22% across Africa as a whole. Given that most Malawi soils cannot supply sufficient Se to crops for adequate human nutrition, the cost and benefits of interventions to alleviate Se deficiency should be determined; for example, Se-enriched nitrogen fertilisers could be adopted as in Finland.
Meekers, Dominique; Van Rossem, Ronan; Silva, Martha; Koleros, Andrew
This study uses data from the 2004 Malawi Demographic and Health Survey to assess the reach of selected radio programs about family planning and health in Malawi and their effect on condom use and discussion of family planning. The results show that such radio programs in Malawi reach a broad audience: eight of the 12 programs were heard by at least half of the respondents, although women were less effectively reached than men. For both women and men, the radio programs were found to have a significant impact on family planning discussion with one's partner. The programs' effect on condom use was limited, however. A positive association was found with ever use of condoms, but no association was found with condom use at last intercourse. This limited impact suggests that such radio communication campaigns need to be informed by research identifying the specific constraints to current condom use in Malawi.
Hurst, Rachel; Siyame, Edwin W. P.; Young, Scott D.; Chilimba, Allan D. C.; Joy, Edward J. M.; Black, Colin R.; Ander, E. Louise; Watts, Michael J.; Chilima, Benson; Gondwe, Jellita; Kang'ombe, Dalitso; Stein, Alexander J.; Fairweather-Tait, Susan J.; Gibson, Rosalind S.; Kalimbira, Alexander A.; Broadley, Martin R.
Selenium (Se) is an essential human micronutrient with critical roles in immune functioning and antioxidant defence. Estimates of dietary Se intakes and status are scarce for Africa although crop surveys indicate deficiency is probably widespread in Malawi. Here we show that Se deficiency is likely endemic in Malawi based on the Se status of adults consuming food from contrasting soil types. These data are consistent with food balance sheets and composition tables revealing that >80% of the Malawi population is at risk of dietary Se inadequacy. Risk of dietary Se inadequacy is >60% in seven other countries in Southern Africa, and 22% across Africa as a whole. Given that most Malawi soils cannot supply sufficient Se to crops for adequate human nutrition, the cost and benefits of interventions to alleviate Se deficiency should be determined; for example, Se-enriched nitrogen fertilisers could be adopted as in Finland. PMID:23478344
Information on the Lake Tahoe watershed, EPA's protection efforts, water quality issues, effects of climate change, Lake Tahoe Total Maximum Daily Load (TMDL), EPA-sponsored projects, list of partner agencies.
Böhne, Astrid; Heule, Corina; Boileau, Nicolas; Salzburger, Walter
Sex determination mechanisms are highly variable across teleost fishes and sexual development is often plastic. Nevertheless, downstream factors establishing the two sexes are presumably conserved. Here, we study sequence evolution and gene expression of core genes of sexual development in a prime model system in evolutionary biology, the East African cichlid fishes. Using the available five cichlid genomes, we test for signs of positive selection in 28 genes including duplicates from the teleost whole-genome duplication, and examine the expression of these candidate genes in three cichlid species. We then focus on a particularly striking case, the A- and B-copies of the aromatase cyp19a1, and detect different evolutionary trajectories: cyp19a1A evolved under strong positive selection, whereas cyp19a1B remained conserved at the protein level, yet is subject to regulatory changes at its transcription start sites. Importantly, we find shifts in gene expression in both copies. Cyp19a1 is considered the most conserved ovary-factor in vertebrates, and in all teleosts investigated so far, cyp19a1A and cyp19a1B are expressed in ovaries and the brain, respectively. This is not the case in cichlids, where we find new expression patterns in two derived lineages: the A-copy gained a novel testis-function in the Ectodine lineage, whereas the B-copy is overexpressed in the testis of the speciest-richest cichlid group, the Haplochromini. This suggests that even key factors of sexual development, including the sex steroid pathway, are not conserved in fish, supporting the idea that flexibility in sexual determination and differentiation may be a driving force of speciation. PMID:23883521
Pariselle, Antoine; Boeger, Walter A.; Snoeks, Jos; Bilong Bilong, Charles F.; Morand, Serge; Vanhove, Maarten P. M.
We discuss geographical distribution and phylogeny of Dactylogyridea (Monogenea) parasitizing Cichlidae to elucidate their hosts' history. Although mesoparasitic Monogenea (Enterogyrus spp.) show typical vicariant distribution, ectoparasitic representatives from different continents are not considered sister taxa, hence their distribution cannot result from vicariance alone. Because of the close host-parasite relationship, this might indicate that present-day cichlid distribution may also reflect dispersal through coastal or brackish waters. Loss of ectoparasites during transoceanic migration, followed by lateral transfer from other fish families might explain extant host-parasite associations. Because of its mesoparasitic nature, hence not subject to salinity variations of the host's environment, Enterogyrus could have survived marine migrations, intolerable for ectoparasites. Host-switches and salinity transitions may be invoked to explain the pattern revealed by a preliminary morphological phylogeny of monogenean genera from Cichlidae and other selected Monogenea genera, rendering the parasite distribution explicable under both vicariance and dispersal. Testable hypotheses are put forward in this parasitological approach to cichlid biogeography. Along with more comprehensive in-depth morphological phylogeny, comparison with molecular data, clarifying dactylogyridean evolution on different continents and from various fish families, and providing temporal information on host-parasite history, are needed to discriminate between the possible scenarios. PMID:21869935
Kerschbaumer, Michaela; Sturmbauer, Christian
Fishes of the family Cichlidae are famous for their spectacular species flocks and therefore constitute a model system for the study of the pathways of adaptive radiation. Their radiation is connected to trophic specialization, manifested in dentition, head morphology, and body shape. Geometric morphometric methods have been established as efficient tools to quantify such differences in overall body shape or in particular morphological structures and meanwhile found wide application in evolutionary biology. As a common feature, these approaches define and analyze coordinates of anatomical landmarks, rather than traditional counts or measurements. Geometric morphometric methods have several merits compared to traditional morphometrics, particularly for the distinction and analysis of closely related entities. Cichlid evolutionary research benefits from the efficiency of data acquisition, the manifold opportunities of analyses, and the potential to visualize shape changes of those landmark-based methods. This paper briefly introduces to the concepts and methods of geometric morphometrics and presents a selection of publications where those techniques have been successfully applied to various aspects of cichlid fish diversification. PMID:21716723
López-Fernández, Hernán; Winemiller, Kirk O.; Montaña, Carmen; Honeycutt, Rodney L.
Genera within the South American cichlid tribe Geophagini display specialized feeding and reproductive strategies, with some taxa specialized for both substrate-sifting and mouth brooding. Several lineages within the clade also possess an epibranchial lobe (EBL), a unique pharyngeal structure that has been proposed to have a function in feeding and/or mouth brooding. A recently published genus-level phylogeny of Neotropical cichlids was used as the evolutionary framework for investigating the evolution of morphological features presumably correlated with diet and mouth brooding in the tribe Geophagini. We tested for possible associations between the geophagine epibranchial lobe and benthic feeding and mouth brooding. We also addressed whether the EBL may be associated with unique patterns of diversification in certain geophagine clades. Tests of binary character correlations revealed the EBL was significantly associated with mouth brooding. We also tested for a relationship between diet and morphology. We analyzed stomach contents and morphometric variation among 21 species, with data for two additional species obtained from the literature. Principal Components Analysis revealed axes of morphological variation significantly correlated with piscivory and benthivory, and both morphology and diet were significantly associated with phylogeny. These results suggest that the EBL could be an adaptation for either feeding or mouth brooding. The EBL, however, was not associated with species richness or accelerated rates of phyletic diversification. PMID:22485154
Bloch, A N; Estela, V J; Leese, J M; Itzkowitz, M
Many monogamous species demonstrate size-assortative mating patterns within natural populations. To better understand the role of intersexual selection in this process, we examined the effect of male preference for female body size in the convict cichlid (Amatitlania siquia). We provided males with a choice between females that differed in size, relative to each other and in relation to the focal male. Based on previous work, we expected males to prefer the largest available female mates across all treatments. Surprisingly, males spent more time near the smaller of two available females, but only when the other female was larger than the male. Additionally, males spent little time with either of two potential female mates when both females were larger than the male. We hypothesized that while males might prefer the largest of available females, female behavior might limit males from acting on this preference. To test this, males were force paired with a smaller or larger female. Pair formation only occurred when the female was smaller than the male, and females that were larger than their male counterparts showed significantly more aggression when compared to smaller females. Together, these data suggest that in the absence of intrasexual competition, male mate preference for large females in convict cichlids might be limited by female aggression.
Egger, Bernd; Klaefiger, Yuri; Theis, Anya; Salzburger, Walter
Although, generally, the origin of sex-limited traits remains elusive, the sensory exploitation hypothesis provides an explanation for the evolution of male sexual signals. Anal fin egg-spots are such a male sexual signal and a key characteristic of the most species-rich group of cichlid fishes, the haplochromines. Males of about 1500 mouth-brooding species utilize these conspicuous egg-dummies during courtship--apparently to attract females and to maximize fertilization success. Here we test the hypothesis that the evolution of haplochromine egg-spots was triggered by a pre-existing bias for eggs or egg-like coloration. To this end, we performed mate-choice experiments in the basal haplochromine Pseudocrenilabrus multicolor, which manifests the plesiomorphic character-state of an egg-spot-less anal fin. Experiments using computer-animated photographs of males indeed revealed that females prefer images of males with virtual ('in-silico') egg-spots over images showing unaltered males. In addition, we tested for color preferences (outside a mating context) in a phylogenetically representative set of East African cichlids. We uncovered a strong preference for yellow, orange or reddish spots in all haplochromines tested and, importantly, also in most other species representing more basal lines. This pre-existing female sensory bias points towards high-quality (carotenoids-enriched) food suggesting that it is adaptive.
Santos, M Emília; Braasch, Ingo; Boileau, Nicolas; Meyer, Britta S; Sauteur, Loïc; Böhne, Astrid; Belting, Heinz-Georg; Affolter, Markus; Salzburger, Walter
The origin of novel phenotypic characters is a key component in organismal diversification; yet, the mechanisms underlying the emergence of such evolutionary novelties are largely unknown. Here we examine the origin of egg-spots, an evolutionary innovation of the most species-rich group of cichlids, the haplochromines, where these conspicuous male fin colour markings are involved in mating. Applying a combination of RNAseq, comparative genomics and functional experiments, we identify two novel pigmentation genes, fhl2a and fhl2b, and show that especially the more rapidly evolving b-paralog is associated with egg-spot formation. We further find that egg-spot bearing haplochromines, but not other cichlids, feature a transposable element in the cis-regulatory region of fhl2b. Using transgenic zebrafish, we finally demonstrate that this region shows specific enhancer activities in iridophores, a type of pigment cells found in egg-spots, suggesting that a cis-regulatory change is causally linked to the gain of expression in egg-spot bearing haplochromines.
Huang, Huateng; Tran, Lucy A P; Knowles, L Lacey
A large number of published phylogenetic estimates are based on a single locus or the concatenation of multiple loci, even though genealogies of single or concatenated loci may not accurately reflect the true history of species diversification (i.e., the species tree). The increased availability of genomic data, coupled with new computational methods, improves resolution of species relationships beyond what was possible in the past. Such developments will no doubt benefit future phylogenetic studies. It remains unclear how robust phylogenies that predate these developments (i.e., the bulk of phylogenetic studies) are to departures from the assumption of strict gene tree-species tree concordance. Here, we present a parametric bootstrap (PBST) approach that assesses the reliability of past phylogenetic estimates in which gene tree-species tree discord was ignored. We focus on a universal cause of discord-the random loss of gene lineages from genetic drift-and apply the method in a meta-analysis of East African cichlids, a group encompassing historical scenarios that are particularly challenging for phylogenetic estimation. Although we identify some evolutionary relationships that are robust to gene tree discord, many past phylogenetic estimates of cichlids are not. We discuss the utility of the PBST method for evaluating the robustness of gene tree-based phylogenetic estimations in general as well as for testing the clade-specific performance of species tree estimation methods and designing sampling strategies that increase the accuracy of estimated species relationships.
Ingraham, Erica; Anderson, Nicole D.; Hurd, Peter L.; Hamilton, Trevor J.
The formation of long-term memories for food sources is essential for the survival of most animals. Long-term memory formation in mammalian species has been demonstrated through a variety of conditioning tasks, however, the nature of long-term memory in fish is less known. In the current study, we explored whether African cichlids (Labidochromis caeruleus) could form memories for food-reinforced stimuli that last for 12 days. During the training sessions, fish were reinforced for approaching an upward drifting line grating. After a rest period of 12 days, fish demonstrated a significant preference for the upward drifting grating. To determine whether this preference could also be reversed, fish were then reinforced for approaching a downward drifting line grating after a 20-day rest period. When tested 12 days later, there were no significant differences in preference for either stimulus; however, following a second training period for the downward stimulus, there was a significant preference for the downward drifting grating. This suggests that cichlids are able to form reversible discrimination-based memories for food-reinforced stimuli that remain consolidated for at least 12 days. PMID:27582695
Muula, Adamson S; Misiri, Humphreys E
Background Physical trauma is an important cause of childhood morbidity and mortality in Africa. There are however, few community-based reports on the subject on the continent. The present study was conducted to explore school children's experience of physical trauma in a disadvantaged periurban area of Blantyre, in Malawi. Methods A cross sectional questionnaire study was carried out among school children in Ndirande-Blantyre, Malawi in 2004. Data were obtained to describe the following aspects of trauma experience: being a victim or observer of motor vehicular accidents involving pedestrians; history of falls from heights; and knowledge about road safety. Sex differences were determined for some of the variables in order to gain insights as to whether there is a difference in trauma experience between boys and girls. Results A total of 217 school children, 99 (45.6%) boys and 118 (54.4%) girls participated in the study. Eight of them reported to have ever been hit by a motor vehicle, 87 (40.1%) had witnessed a road accident where a pedestrian had been hit and 83 (38.2%) had witnessed a pedestrian they knew having been hit by a motor vehicle. Of those that reported to have ever been hit by motor vehicle, 2 (25%) reported that they had been hospitalized as a result of injury. With regard to falling from heights, 86 reported to have ever fallen from tree, 44 of these (51.2%) were injured from the fall and 14 (16.3%) were hospitalized as a result of injury sustained from the fall. Girls were more likely to fall from trees and getting injured as compared to males (p = 0.04 for both situations). Just under half (41.9%) of the study participants were able to report the correct procedure of crossing the road despite the fact that the majority (80%) reported having been taught road safety at home or school. Conclusion Many school children in Blantyre, Malawi have been exposed to trauma either involving themselves or someone they observed. Prevention, including education
"Despite increasing prominence in international population movements, refugee migration is a poorly understood social and spatial process. Common assumptions about refugee migration have rarely been empirically tested. This article focuses attention on the migration decision-making process of refugees. Its conceptual basis is that the decision making of refugees can be critically compared with that of other migrants. An analytical framework is suggested which focuses on the receipt and evaluation of information about prospective destinations in the decision-making process. It is tested in the context of Mozambican refugees in Malawi in 1992 and 1993. Analysis shows that this framework achieves a qualified success in explaining the patterns and processes of migration of the study population, and suggests broader applications for the framework."
Koser (1993) proposed a research model for examining the interaction between information and repatriation, which focused upon the receipt and evaluation of information about their homes by refugees in exile. That model is presented and tested using interview responses from Mozambican refugees in Malawi sampled in 1992. The goal was to disaggregate refugees' decision-making process in an attempt to understand the repatriation process. Findings are based upon the interview responses of 125 randomly selected refugee household heads, all members of 50 of the households, and local leaders. The study found that such a narrow focus will fail to yield any comprehensive explanation. However, the specific focus upon information was determined to have certain empirical, conceptual, and policy-oriented advantages.
Numerous studies have documented the determinants of sexual behavior among adolescents in less-developed countries, yet relatively little is known about the influence of social contexts such as school and neighborhood. Using two waves of data from a school-based longitudinal survey conducted in Malawi from 2011-13, this study advances our understanding of the relationship between school-level socioeconomic contexts and adolescents' sexual activity. The results from two-level multinomial logistic regression models suggest that high socioeconomic composition of the student body in school decreases the odds of initiation of sexual activity, independent of other important features of schools and individual-level characteristics. This study also finds that the association between school socioeconomic composition and sexual activity is statistically significant among male adolescents but not female adolescents, suggesting that schools' socioeconomic contexts may be more relevant to male adolescents' initiation of sexual activity.
Background Maternal mortality, although largely preventable, remains unacceptably high in developing countries such as Malawi and creates a number of intergenerational impacts. Few studies have investigated the far-reaching impacts of maternal death beyond infant survival. This study demonstrates the short- and long-term impacts of maternal death on children, families, and the community in order to raise awareness of the true costs of maternal mortality and poor maternal health care in Neno, a rural and remote district in Malawi. Methods Qualitative in-depth interviews were conducted to assess the impact of maternal mortality on child, family, and community well-being. We conducted 20 key informant interviews, 20 stakeholder interviews, and six sex-stratified focus group discussions in the seven health centers that cover the district. Transcripts were translated, coded, and analyzed in NVivo 10. Results Participants noted a number of far-reaching impacts on orphaned children, their new caretakers, and extended families following a maternal death. Female relatives typically took on caregiving responsibilities for orphaned children, regardless of the accompanying financial hardship and frequent lack of familial or governmental support. Maternal death exacerbated children’s vulnerabilities to long-term health and social impacts related to nutrition, education, employment, early partnership, pregnancy, and caretaking. Impacts were particularly salient for female children who were often forced to take on the majority of the household responsibilities. Participants cited a number of barriers to accessing quality child health care or support services, and many were unaware of programming available to assist them in raising orphaned children or how to access these services. Conclusions In order to both reduce preventable maternal mortality and diminish the impacts on children, extended families, and communities, our findings highlight the importance of financing and
Hoegg, Simone; Boore, Jeffrey L; Kuehl, Jennifer V; Meyer, Axel
Background Teleost fish have seven paralogous clusters of Hox genes stemming from two complete genome duplications early in vertebrate evolution, and an additional genome duplication during the evolution of ray-finned fish, followed by the secondary loss of one cluster. Gene duplications on the one hand, and the evolution of regulatory sequences on the other, are thought to be among the most important mechanisms for the evolution of new gene functions. Cichlid fish, the largest family of vertebrates with about 2500 species, are famous examples of speciation and morphological diversity. Since this diversity could be based on regulatory changes, we chose to study the coding as well as putative regulatory regions of their Hox clusters within a comparative genomic framework. Results We sequenced and characterized all seven Hox clusters of Astatotilapia burtoni, a haplochromine cichlid fish. Comparative analyses with data from other teleost fish such as zebrafish, two species of pufferfish, stickleback and medaka were performed. We traced losses of genes and microRNAs of Hox clusters, the medaka lineage seems to have lost more microRNAs than the other fish lineages. We found that each teleost genome studied so far has a unique set of Hox genes. The hoxb7a gene was lost independently several times during teleost evolution, the most recent event being within the radiation of East African cichlid fish. The conserved non-coding sequences (CNS) encompass a surprisingly large part of the clusters, especially in the HoxAa, HoxCa, and HoxDa clusters. Across all clusters, we observe a trend towards an increased content of CNS towards the anterior end. Conclusion The gene content of Hox clusters in teleost fishes is more variable than expected, with each species studied so far having a different set. Although the highest loss rate of Hox genes occurred immediately after whole genome duplications, our analyses showed that gene loss continued and is still ongoing in all teleost
Background Among multigene families, ribosomal RNA (rRNA) genes are the most frequently studied and have been explored as cytogenetic markers to study the evolutionary history of karyotypes among animals and plants. In this report, we applied cytogenetic and genomic methods to investigate the organization of rRNA genes among cichlid fishes. Cichlids are a group of fishes that are of increasing scientific interest due to their rapid and convergent adaptive radiation, which has led to extensive ecological diversity. Results The present paper reports the cytogenetic mapping of the 5S rRNA genes from 18 South American, 22 African and one Asian species and the 18S rRNA genes from 3 African species. The data obtained were comparatively analyzed with previously published information related to the mapping of rRNA genes in cichlids. The number of 5S rRNA clusters per diploid genome ranged from 2 to 15, with the most common pattern being the presence of 2 chromosomes bearing a 5S rDNA cluster. Regarding 18S rDNA mapping, the number of sites ranged from 2 to 6, with the most common pattern being the presence of 2 sites per diploid genome. Furthermore, searching the Oreochromis niloticus genome database led to the identification of a total of 59 copies of 5S rRNA and 38 copies of 18S rRNA genes that were distributed in several genomic scaffolds. The rRNA genes were frequently flanked by transposable elements (TEs) and spread throughout the genome, complementing the FISH analysis that detect only clustered copies of rRNA genes. Conclusions The organization of rRNA gene clusters seems to reflect their intense and particular evolutionary pathway and not the evolutionary history of the associated taxa. The possible role of TEs as one source of rRNA gene movement, that could generates the spreading of ribosomal clusters/copies, is discussed. The present paper reinforces the notion that the integration of cytogenetic data and genomic analysis provides a more complete picture for
McDonnell, Laura H.; Chapman, Lauren J.
Tropical inland fishes are predicted to be especially vulnerable to thermal stress because they experience small temperature fluctuations that may select for narrow thermal windows. In this study, we measured resting metabolic rate (RMR), critical oxygen tension (Pcrit) and critical thermal maximum (CTMax) of the widespread African cichlid (Pseudocrenilabrus multicolor victoriae) in response to short-term acclimation to temperatures within and above their natural thermal range. Pseudocrenilabrus multicolor collected in Lake Kayanja, Uganda, a population living near the upper thermal range of the species, were acclimated to 23, 26, 29 and 32°C for 3 days directly after capture, and RMR and Pcrit were then quantified. In a second group of P. multicolor from the same population, CTMax and the thermal onset of agitation were determined for fish acclimated to 26, 29 and 32°C for 7 days. Both RMR and Pcrit were significantly higher in fish acclimated to 32°C, indicating decreased tolerance to hypoxia and increased metabolic requirements at temperatures only slightly (∼1°C) above their natural thermal range. The CTMax increased with acclimation temperature, indicating some degree of thermal compensation induced by short-term exposure to higher temperatures. However, agitation temperature (likely to represent an avoidance response to increased temperature during CTMax trials) showed no increase with acclimation temperature. Overall, the results of this study demonstrate that P. multicolor is able to maintain its RMR and Pcrit across the range of temperatures characteristic of its natural habitat, but incurs a higher cost of resting metabolism and reduced hypoxia tolerance at temperatures slightly above its present range. PMID:27293734
Simões, José M; Teles, Magda C; Oliveira, Rui F; Van der Linden, Annemie; Verhoye, Marleen
The African cichlid Oreochromis mossambicus (Mozambique tilapia) has been used as a model system in a wide range of behavioural and neurobiological studies. The increasing number of genetic tools available for this species, together with the emerging interest in its use for neurobiological studies, increased the need for an accurate hodological mapping of the tilapia brain to supplement the available histological data. The goal of our study was to elaborate a three-dimensional, high-resolution digital atlas using magnetic resonance imaging, supported by Nissl staining. Resulting images were viewed and analysed in all orientations (transverse, sagittal, and horizontal) and manually labelled to reveal structures in the olfactory bulb, telencephalon, diencephalon, optic tectum, and cerebellum. This high resolution tilapia brain atlas is expected to become a very useful tool for neuroscientists using this fish model and will certainly expand their use in future studies regarding the central nervous system.
Peeke, H V; Avis, H H; Peeke, S C
Four experiments using territorial Convict Cichlids investigated motivational factors involved in the incremental and decremental processes associated with aggression resulting from exposure to conspecifics intruded into the territory. The first three experiments varied some single aspect of the experimental situation (temperature, distance from the nest or size of the intruder). The fourth experiment combined those factors which resulted in faster habituation (small intruder, far from the nest, in cool water) and compared the response to factors which resulted in slower habituation or an increase in response rate (large intruder, close to the nest in warm water). While a combination of higher intensity stimuli did result in slower habituation than the combination of lower intensity stimuli, response rate was not a simple algebraic summation of the factors. Results are discussed in relation to multi-factor theory of habituation and the nature of "drive".
Simões, José M.; Teles, Magda C.; Oliveira, Rui F.; Van der Linden, Annemie; Verhoye, Marleen
The African cichlid Oreochromis mossambicus (Mozambique tilapia) has been used as a model system in a wide range of behavioural and neurobiological studies. The increasing number of genetic tools available for this species, together with the emerging interest in its use for neurobiological studies, increased the need for an accurate hodological mapping of the tilapia brain to supplement the available histological data. The goal of our study was to elaborate a three-dimensional, high-resolution digital atlas using magnetic resonance imaging, supported by Nissl staining. Resulting images were viewed and analysed in all orientations (transverse, sagittal, and horizontal) and manually labelled to reveal structures in the olfactory bulb, telencephalon, diencephalon, optic tectum, and cerebellum. This high resolution tilapia brain atlas is expected to become a very useful tool for neuroscientists using this fish model and will certainly expand their use in future studies regarding the central nervous system. PMID:22984463
Ma, Yunyong; Hu, Caroline K.; Huguenard, John R.; Fernald, Russell D.
Initiating and regulating vertebrate reproduction requires pulsatile release of gonadotropin-releasing hormone (GnRH1) from the hypothalamus. Coordinated GnRH1 release, not simply elevated absolute levels, effects the release of pituitary gonadotropins that drive steroid production in the gonads. However, the mechanisms underlying synchronization of GnRH1 neurons are unknown. Control of synchronicity by gap junctions between GnRH1 neurons has been proposed but not previously found. We recorded simultaneously from pairs of transgenically labeled GnRH1 neurons in adult male Astatotilapia burtoni cichlid fish. We report that GnRH1 neurons are strongly and uniformly interconnected by electrical synapses that can drive spiking in connected cells and can be reversibly blocked by meclofenamic acid. Our results suggest that electrical synapses could promote coordinated spike firing in a cellular assemblage of GnRH1 neurons to produce the pulsatile output necessary for activation of the pituitary and reproduction. PMID:25775522
Sookrajh, Reshma; McCloud, Salanjira
We review spaces of contest in the formulation and implementation of the Religious Education curriculum through an exploration of the understandings regarding the dual-mode Religious Education (RE) curriculum policy for secondary schools in Malawi. Six leaders of nationally organised faith communities in Malawi were interviewed about their…
Schweitzer, Cécile; Melot, Geoffrey; Laubu, Chloé; Teixeira, Maria; Motreuil, Sébastien; Dechaume-Moncharmont, François-Xavier
In monogamous species, partner compatibility is a key factor influencing pairing and reproductive success. In pairs with biparental care, studies have mostly focused on behavioral compatibility because it is likely to encourage the coordination of parental care within pairs, leading to a better reproductive success. Behavior modulation, throughout the breeding season and as a function of the social context, is under the regulatory feedback control of endocrine mechanisms. From this link, the idea of hormonal partner compatibility as a key component of pair cohesion and maintenance has recently emerged. Here, we investigated the link between partner behavioral assortment and their hormonal response to the pairing context. We formed assortative and disassortative pairs of convict cichlids based on their behavioral type (proactive or reactive) and took hormone and fitness measurements. Testosterone, 11-ketotestosterone, 17β-estradiol and cortisol levels were measured from fish-holding water before and after pair formation. We found no relationship between the behavioral type of individuals and their pre-pairing hormone levels. Only the level of cortisol was affected by the partner but independently of its behavioral type. Reproductive success was not affected by the level of hormonal similarity within pairs, but we found that the variation in 11-ketotestosterone similarity between the isolated context and the pairing context was related to spawning size, and the variation in cortisol similarity to the number of fry. Behavioral compatibility does not reflect hormonal compatibility in the convict cichlid, but the relationship between reproductive success and the flexibility of hormonal similarity between partners suggests hormonal adjustment within pairs in this species.
Almeida, Olinda; Oliveira, Rui F
The nonapeptide arginine vasotocin (AVT) and its mammalian homologue arginine vasopressin play a key role in the regulation of social behaviour across vertebrates. In teleost fishes, three AVT neuronal populations have been described in the preoptic area (POA): the parvocellular (pPOA), the magnocellular (mPOA) and the gigantocellular (gPOA). Neurons from each of these areas project both to the pituitary and to other brain regions, where AVT is supposed to regulate neural circuits underlying social behaviour. However, in the fish species studied so far, there is considerable variation in which AVT neuronal populations are involved in behavioural modulation and in the direction of the effect. In this study, the association between AVT neuronal phenotypes and social status was investigated in the Mozambique tilapia (Oreochromis mossambicus). This species is an African female mouth-brooding cichlid fish in which males form breeding aggregations in which dominant males establish territories and subordinate males to act as floaters. With respect to sex differences in AVT neuronal phenotypes, females have a larger number of AVT neurons in the pPOA and mPOA. Within males, AVT appeared associated with social subordination, as indicated by the larger cell body areas of AVT neurons in mPOA and gPOA nuclei of non-territorial males. There were also positive correlations between submissive behaviour and the soma size of AVT cells in all three nuclei and AVT cell number in the mPOA. In summary, the results provide evidence for an involvement of AVT in the modulation of social behaviour in tilapia, but it was not possible to identify specific roles for specific AVT neuronal populations. The results presented here also contrast with those previously published for another cichlid species with a similar mating system, which highlights the species-specific nature of the pattern of association between AVT and social behaviour even within the same taxonomic family.
Vanhove, Maarten P M; Van Steenberge, Maarten; Dessein, Steven; Volckaert, Filip A M; Snoeks, Jos; Huyse, Tine; Pariselle, Antoine
Fishes normally restricted to inland waters are valuable model systems for historical biogeography, inter alia, because of their limited dispersal abilities and concordance with the distribution patterns of other freshwater taxa (Zogaris et al. 2009). The comparison of fish species assemblages has been the major biogeographical tool for delineating African aquatic ecoregions as the fossil record is often meagre and merely offers complementary information. This is, for example, the case for the Zambezian and Congolian ichthyofaunal provinces, which display substantial contemporary fish diversity (Stewart 2001). Between both regions lies the Bangweulu-Mweru ecoregion (sensu Scott 2005), known for its high percentage of endemicity. Although hydrographically belonging to the Congo Basin, the Bangweulu-Mweru ecoregion has a high affinity with the Zambezi province (Scott 2005), due to historical river connections (Tweddle 2010). Studies comparing the Zambezi and Congo ichthyofaunal provinces are rare and hampered by lack of data from the Congo Basin. The latter harbours more than 1250 fish species (Snoeks et al. 2011) while in the Zambezi, only 120 freshwater fishes are found (Tweddle 2010). Indeed, species richness declines in all major African teleost families from the Congo Basin southwards, riverine haplochromine cichlids forming a notable exception to this rule (Joyce et al. 2005). Although it was hypothesized by Tweddle (2010) that the origin of many Zambezian fish species is in the Congo Basin, the haplochromines Serranochromis Regan, Sargochromis Regan, Pharyngochromis Greenwood and Chetia Trewavas, together forming the serranochromines, have their centre of diversity in the rivers of the Zambezian ichthyofaunal province (Joyce et al. 2005). Therefore, the biogeographical history of Cichlidae across the Zambezi- Congo watershed is not only key to cichlid biogeography on an African scale, but also complementary to biogeography of all other teleosts in the region
Theis, Anya; Salzburger, Walter; Egger, Bernd
Color and pigmentation patterns of animals are often targets of sexual selection because of their role in communication. Although conspicuous male traits are typically implicated with intersexual selection, there are examples where sex-specific displays play a role in an intrasexual context, e.g. when they serve as signals for aggression level and/or status. Here, we focus on the function of a conspicuous male ornament in the most species-rich tribe of cichlid fishes, the haplochromines. A characteristic feature of these ca. 1500 species are so-called egg-spots in form of ovoid markings on the anal fins of males, which are made up of carotenoid based pigment cells. It has long been assumed that these yellow, orange or reddish egg-spots play an important role in the courtship and spawning behavior of these maternal mouth-brooding fishes by mimicking the eggs of a conspecific female. The exact function of egg-spots remains unknown, however, and there are several hypotheses about their mode of action. To uncover the function of this cichlid-specific male ornament, we used female mate choice experiments and a male aggression test in the haplochromine species Astatotilapia burtoni. We manipulated the number and arrangement of egg-spots on the anal fins of males, or removed them entirely, and tested (1) female preference with visual contact only using egg-traps, (2) female preference with free contact using paternity testing with microsatellites and (3) male aggression. We found that females did not prefer males with many egg-spots over males with fewer egg-spots and that females tended to prefer males without egg-spots over males with egg-spots. Importantly, males without egg-spots sired clutches with the same fertilization rate as males with egg-spots. In male aggression trials, however, males with fewer egg-spots received significantly more attacks, suggesting that egg-spots are an important signal in intrasexual communication.
Lupafya, Phindile Chitsulo; Mwagomba, Beatrice L Matanje; Hosig, Kathy; Maseko, Lucy M; Chimbali, Henry
Malawi is a Sub-Saharan African country experiencing the epidemiological transition from predominantly infectious to noncommunicable diseases (NCDs) with dramatically increasing prevalence of lifestyle-related diseases such as obesity, hypertension, and diabetes. Malawi's 2011-2016 Health Sector Strategic Plan included NCDs, and an NCD Control Program was established with subsequent development of a National Action Plan for prevention and management of NCDs launched in 2013. The current study was designed to identify gaps in implementation of NCD control program policies and action plan strategies by describing current efforts toward prevention and management of NCDs in Malawi with emphasis on challenges and opportunities. Semistructured questionnaires were used to collect quantitative and qualitative data from Malawi Ministry of Health personnel (senior officers, service providers, health education officers, and nutritionists) in 10 health districts and 3 central hospitals. Frequencies were generated for quantitative data. Qualitative data were used to generate themes and most common responses. Results showed that current services focus on facility-based NCD screening and clinical services rather than active screening, prevention, and community awareness and outreach, although respondents emphasized the importance of prevention, lifestyle education, and community outreach. Respondents indicated inadequate resources for NCD services including financial capital, human resources, equipment and supplies, and transportation. While Malawi has begun to address NCDs, policy and practice implications include (a) better integration of services within the existing infrastructure with emphasis on capacity building; (b) greater implementation of planned NCD activities; (c) a stronger, more comprehensive data management system; and (d) innovative funding solutions.
Maruska, Karen P; Ung, Uyhun S; Fernald, Russell D
Sexual reproduction in all animals depends on effective communication between signalers and receivers. Many fish species, especially the African cichlids, are well known for their bright coloration and the importance of visual signaling during courtship and mate choice, but little is known about what role acoustic communication plays during mating and how it contributes to sexual selection in this phenotypically diverse group of vertebrates. Here we examined acoustic communication during reproduction in the social cichlid fish, Astatotilapia burtoni. We characterized the sounds and associated behaviors produced by dominant males during courtship, tested for differences in hearing ability associated with female reproductive state and male social status, and then tested the hypothesis that female mate preference is influenced by male sound production. We show that dominant males produce intentional courtship sounds in close proximity to females, and that sounds are spectrally similar to their hearing abilities. Females were 2-5-fold more sensitive to low frequency sounds in the spectral range of male courtship sounds when they were sexually-receptive compared to during the mouthbrooding parental phase. Hearing thresholds were also negatively correlated with circulating sex-steroid levels in females but positively correlated in males, suggesting a potential role for steroids in reproductive-state auditory plasticity. Behavioral experiments showed that receptive females preferred to affiliate with males that were associated with playback of courtship sounds compared to noise controls, indicating that acoustic information is likely important for female mate choice. These data show for the first time in a Tanganyikan cichlid that acoustic communication is important during reproduction as part of a multimodal signaling repertoire, and that perception of auditory information changes depending on the animal's internal physiological state. Our results highlight the
Background Cichlid fishes have undergone rapid, expansive evolutionary radiations that are manifested in the diversification of their trophic morphologies, tooth patterning and coloration. Understanding the molecular mechanisms that underlie the cichlids' unique patterns of evolution requires a thorough examination of genes that pattern the neural crest, from which these diverse phenotypes are derived. Among those genes, the homeobox-containing Dlx gene family is of particular interest since it is involved in the patterning of the brain, jaws and teeth. Results In this study, we characterized the dlx genes of an African cichlid fish, Astatotilapia burtoni, to provide a baseline to later allow cross-species comparison within Cichlidae. We identified seven dlx paralogs (dlx1a, -2a, -4a, -3b, -4b, -5a and -6a), whose orthologies were validated with molecular phylogenetic trees. The intergenic regions of three dlx gene clusters (dlx1a-2a, dlx3b-4b, and dlx5a-6a) were amplified with long PCR. Intensive cross-species comparison revealed a number of conserved non-coding elements (CNEs) that are shared with other percomorph fishes. This analysis highlighted additional lineage-specific gains/losses of CNEs in different teleost fish lineages and a novel CNE that had previously not been identified. Our gene expression analyses revealed overlapping but distinct expression of dlx orthologs in the developing brain and pharyngeal arches. Notably, four of the seven A. burtoni dlx genes, dlx2a, dlx3b, dlx4a and dlx5a, were expressed in the developing pharyngeal teeth. Conclusion This comparative study of the dlx genes of A. burtoni has deepened our knowledge of the diversity of the Dlx gene family, in terms of gene repertoire, expression patterns and non-coding elements. We have identified possible cichlid lineage-specific changes, including losses of a subset of dlx expression domains in the pharyngeal teeth, which will be the targets of future functional studies. PMID:21205289
tkint, Tim; Verheyen, Erik; De Kegel, Barbara; Helsen, Philippe; Adriaens, Dominique
Background As in any vertebrate, heads of fishes are densely packed with functions. These functions often impose conflicting mechanical demands resulting in trade-offs in the species-specific phenotype. When phenotypical traits are linked to gender-specific parental behavior, we expect sexual differences in these trade-offs. This study aims to use mouthbrooding cichlids as an example to test hypotheses on evolutionary trade-offs between intricately linked traits that affect different aspects of fitness. We focused on the oral apparatus, which is not only equipped with features used to feed and breathe, but is also used for the incubation of eggs. We used this approach to study mouthbrooding as part of an integrated functional system with diverging performance requirements and to explore gender-specific selective environments within a species. Methodology/Principal Findings Because cichlids are morphologically very diverse, we hypothesize that the implications of the added constraint of mouthbrooding will primarily depend on the dominant mode of feeding of the studied species. To test this, we compared the trade-off for two maternal mouthbrooding cichlid species: a “suction feeder” (Haplochromis piceatus) and a “biter” (H. fischeri). The comparison of morphology and performance of both species revealed clear interspecific and intersex differences. Our observation that females have larger heads was interpreted as a possible consequence of the fact that in both the studied species mouthbrooding is done by females only. As hypothesized, the observed sexual dimorphism in head shape is inferred as being suboptimal for some aspects of the feeding performance in each of the studied species. Our comparison also demonstrated that the suction feeding species had smaller egg clutches and more elongated eggs. Conclusions/Significance Our findings support the hypothesis that there is a trade-off between mouthbrooding and feeding performance in the two studied
Rehage, Jennifer S.; Dunlop, Katherine L.; Loftus, William F.
The strong impact of non-native predators in aquatic systems is thought to relate to the evolutionary naiveté of prey. Due to isolation and limited dispersal, this naiveté may be relatively high in freshwater systems. In this study, we tested this notion by examining the antipredator response of native mosquitofish, Gambusia holbrooki, to two non-native predators found in the Everglades, the African jewelfish,Hemichromis letourneuxi, and the Mayan cichlid, Cichlasoma urophthalmus. We manipulated prey naiveté by using two mosquitofish populations that varied in their experience with the recent invader, the African jewelfish, but had similar levels of experience with the longer-established Mayan cichlid. Specifically, we tested these predictions: (1) predator hunting modes differed between the two predators, (2) predation rates would be higher by the novel jewelfish predator, (3) particularly on the naive population living where jewelfish have not invaded yet, (4) antipredator responses would be stronger to Mayan cichlids due to greater experience and weaker and/or ineffective to jewelfish, and (5) especially weakest by the naive population. We assayed prey and predator behavior, and prey mortality in lab aquaria where both predators and prey were free-ranging. Predator hunting modes and habitat domains differed, with jewelfish being more active search predators that used slightly higher parts of the water column and less of the habitat structure relative to Mayan cichlids. In disagreement with our predictions, predation rates were similar between the two predators, antipredator responses were stronger to African jewelfish (except for predator inspections), and there was no difference in response between jewelfish-savvy and jewelfish-naive populations. These results suggest that despite the novelty of introduced predators, prey may be able to respond appropriately if non-native predator archetypes are similar enough to those of native predators, if prey rely
Snekser, Jennifer L.; Itzkowitz, Murray
The convict cichlid (Amatitlania nigrofasciata) has been extensively examined in relation to many behavioral topics, such as courtship, pair-bonding, bi-parental care, and territoriality. Recently, this model species has been utilized in studies on genetics, endocrinology, and neuroanatomy, with an ultimate goal of connecting behavior with its underlying mechanisms. The goal of this study was two-fold: (1) profile the circulating levels of plasma 11KT in the male convict cichlid at multiple points during the reproductive cycle and (2) generally compare the hormonal profiles of the widely used laboratory populations and those of a free-living population in the streams of Costa Rica. The results of the field experiment showed that male convict cichlids had higher levels of circulating 11KT during courtship and lower during the parental care and non-breeding phases. The profile of the laboratory population was similar to the profile of the free-living individuals, with significantly higher levels of 11KT occurring during courtship than during parental care, though the level of 11KT during non-breeding phase was elevated in the laboratory. The high levels of 11KT during courtship and low levels of 11KT during parental care found in both the field and the laboratory is similar to what has been reported in other species of teleosts, and may suggest an important function of 11KT in the expression of courtship behavior and the subsequent onset of parental behaviors in this model species. PMID:26020006
Valente, G Targino; Vitorino, C de Andrade; Cabral-de-Mello, D C; Oliveira, C; Souza, I Lima; Martins, C; Venere, P C
Cichlids represent one of the most species-rich families of fishes and have attracted the attention of evolutionary biologists due to the rapid radiation occurring in some groups and the importance of some species in the world aquaculture. Cytogenetic analysis was conducted in 10 cichlid species from the Araguaia River, Amazon Basin, Brazil. The chromosome number was 2n=48 for all analyzed species except for Laetacara araguaiae Ottoni et Costa, 2009 (2n=44). Chromosomal polymorphism was detected only in Geophagus proximus (Castelnau, 1855), which exhibits an extra large submetacentric and and a dot-like chromosomes. Moreover, the C-banding revealed a general pericentromeric heterochromatic pattern and some additional blocks for some species. The heterochromatic blocks corresponding to AgNOR bearing regions were observed in all species and also corresponded to CMA3 positive blocks, which were observed in terminal regions. Besides the general conserved chromosomal and heterochromatin patterns for South American cichlids, the presence of GC-rich heterochromatin was quite different in the species Biotodoma cupido (Heckel, 1840), Geophagus proximus, Retroculus lapidifer (Castelnau, 1855), Crenicichla strigata Günther, 1862 and Heros efasciatus Heckel, 1840. The results suggest that independent events of heterochromatin modification occurred during chromosome evolution in the group, regardless of the conservation of macro-chromosomal structure.
Gunter, Helen M; Fan, Shaohua; Xiong, Fan; Franchini, Paolo; Fruciano, Carmelo; Meyer, Axel
Adaptive phenotypic plasticity, the ability of an organism to change its phenotype to match local environments, is increasingly recognized for its contribution to evolution. However, few empirical studies have explored the molecular basis of plastic traits. The East African cichlid fish Astatoreochromis alluaudi displays adaptive phenotypic plasticity in its pharyngeal jaw apparatus, a structure that is widely seen as an evolutionary key innovation that has contributed to the remarkable diversity of cichlid fishes. It has previously been shown that in response to different diets, the pharyngeal jaws change their size, shape and dentition: hard diets induce an adaptive robust molariform tooth phenotype with short jaws and strong internal bone structures, while soft diets induce a gracile papilliform tooth phenotype with elongated jaws and slender internal bone structures. To gain insight into the molecular underpinnings of these adaptations and enable future investigations of the role that phenotypic plasticity plays during the formation of adaptive radiations, the transcriptomes of the two divergent jaw phenotypes were examined. Our study identified a total of 187 genes whose expression differs in response to hard and soft diets, including immediate early genes, extracellular matrix genes and inflammatory factors. Transcriptome results are interpreted in light of expression of candidate genes-markers for tooth size and shape, bone cells and mechanically sensitive pathways. This study opens up new avenues of research at new levels of biological organization into the roles of phenotypic plasticity during speciation and radiation of cichlid fishes.
Arbour, J H; López-Fernández, H
Morphological, lineage and ecological diversity can vary substantially even among closely related lineages. Factors that influence morphological diversification, especially in functionally relevant traits, can help to explain the modern distribution of disparity across phylogenies and communities. Multivariate axes of feeding functional morphology from 75 species of Neotropical cichlid and a stepwise-AIC algorithm were used to estimate the adaptive landscape of functional morphospace in Cichlinae. Adaptive landscape complexity and convergence, as well as the functional diversity of Cichlinae, were compared with expectations under null evolutionary models. Neotropical cichlid feeding function varied primarily between traits associated with ram feeding vs. suction feeding/biting and secondarily with oral jaw muscle size and pharyngeal crushing capacity. The number of changes in selective regimes and the amount of convergence between lineages was higher than expected under a null model of evolution, but convergence was not higher than expected under a similarly complex adaptive landscape. Functional disparity was compatible with an adaptive landscape model, whereas the distribution of evolutionary change through morphospace corresponded with a process of evolution towards a single adaptive peak. The continentally distributed Neotropical cichlids have evolved relatively rapidly towards a number of adaptive peaks in functional trait space. Selection in Cichlinae functional morphospace is more complex than expected under null evolutionary models. The complexity of selective constraints in feeding morphology has likely been a significant contributor to the diversity of feeding ecology in this clade.
... Swiss shores of Lake Constance at the town of Rorschach. Eutrophication, or the process of nutrient enrichment, is rapidly accelerated ... of the value of Lake Constance, efforts to mitigate eutrophication were initiated in the 1970's. MISR was built and is managed ...
Background Malawi faces a severe shortage of health workers, a factor that has contributed greatly to high maternal mortality in the country. Most clinical care is performed by mid-level providers (MLPs). While utilization of these cadres in providing health care is a solution to the current shortages, demotivating factors within the Malawian health system are pushing them into private, non-governmental, and other non-health related positions. This study aims to highlight these demotivating factors by exploring the critical aspects that influence MLPs’ intention to leave their jobs. Methods This descriptive qualitative study formed part of the larger Health Systems Strengthening for Equity (HSSE) study. Data presented in this paper were collected in Malawi using the Critical Incident Analysis tool. Participants were asked to narrate an incident that had happened during the past three months which had made them seriously consider leaving their job. Data were subjected to thematic analysis using NVivo 8 software. Results Of the 84 respondents who participated in a Critical Incident Analysis interview, 58 respondents (69%) indicated they had experienced a demotivating incident in the previous three months that had made them seriously consider leaving their job. The most commonly cited critical factors were being treated unfairly or with disrespect, lack of recognition of their efforts, delays and inconsistencies in salary payments, lack of transparent processes and criteria for upgrading or promotion, and death of patients. Conclusion Staff motivation and an enabling environment are crucial factors for retaining MLPs in the Malawian health system. This study revealed key ‘tipping points’ that drive staff to seriously consider leaving their jobs. Many of the factors underlying these critical incidents can be addressed by improved management practices and the introduction of fair and transparent policies. Managers need to be trained and equipped with effective
Holzman, Roi; Hulsey, C. Darrin
Hybrid phenotypes are often intermediate between those of parental species. However, hybridization can generate novel phenotypes when traits are complex. For instance, even when the morphologies of individual musculo-skeletal components do not segregate outside the parental range in hybrid offspring, complex functional systems can exhibit emergent phenotypes whose mechanics exceed the parental values. To determine if transgression in mechanics could facilitate divergence during an adaptive radiation, we examined three functional systems in the trophic apparatus of Lake Malawi cichlid fishes. We conducted a simulation study of hybridization between species pairs whose morphology for three functional systems was empirically measured, to determine how the evolutionary divergence of parental species influences the frequency that hybridization could produce mechanics that transgress the parental range. Our simulations suggest that the complex mechanical systems of the cichlid trophic apparatus commonly exhibit greater transgression between more recently diverged cichlid species. Because (1) all three mechanical systems produce hybrids with transgressive mechanics in Lake Malawi cichlids, (2) hybridization is common, and (3) single hybrid crosses often recapitulate a substantial diversity of mechanics, we conclude that mechanical transgressive segregation could play an important role in the rapid accumulation of phenotypic variation in adaptive radiations. PMID:28079133
Mtambo, Owen P L; Masangwi, Salule J; Kazembe, Lawrence N M
Analyses of childhood overweight have mainly used mean regression. However, using quantile regression is more appropriate as it provides flexibility to analyse the determinants of overweight corresponding to quantiles of interest. The main objective of this study was to fit a Bayesian additive quantile regression model with structured spatial effects for childhood overweight in Malawi using the 2010 Malawi DHS data. Inference was fully Bayesian using R-INLA package. The significant determinants of childhood overweight ranged from socio-demographic factors such as type of residence to child and maternal factors such as child age and maternal BMI. We observed significant positive structured spatial effects on childhood overweight in some districts of Malawi. We recommended that the childhood malnutrition policy makers should consider timely interventions based on risk factors as identified in this paper including spatial targets of interventions.
Although female sterilization is the most widely used modern contraceptive method in the world, most family planning programs in Africa have had difficulty providing it. Malawi, however, despite daunting constraints, has made female sterilization widely and equitably accessible, thereby increasing method choice and helping its citizens better meet their reproductive intentions. Ten percent of currently married Malawian women of reproductive age rely on female sterilization for contraceptive protection, compared with less than 2 percent across Africa, and demand to limit births now exceeds demand to space births. Malawi's female sterilization prevalence surpasses that of some high-resource countries. Key service-delivery factors enabling this achievement include supportive policies, strong public-private partnerships, and mobile services delivered at no cost by dedicated providers. Challenges remain, but Malawi's achievement offers lessons for other countries with low availability of female sterilization and similar resource constraints.
Chilemba, Winnie; van Wyk, Neltjie; Leech, Ronell
Our aim for this study was to describe emotional abuse as it is experienced by women living with HIV and AIDS in Malawi. The study was conducted in the Lilongwe district in Malawi and used a descriptive phenomenological approach. Twelve women from two public health care clinics under the Lilongwe District Health Office were interviewed. Violating experiences that scarred the personhood and inherent value of being human were found to be the essence of their emotional abuse. Their husbands, family, and community members were responsible for the humiliation, abandonment, and blaming that caused them to feel hopeless.
Climate change poses a big challenge to rural livelihoods in the Shire Valley area of Southern Malawi, where communities have depended almost entirely on rain-fed agriculture for generations. The Shire Valley area comprises of low-altitude dambo areas and uplands which have been the main agricultural areas. Since early to mid 1980s, the uplands have experienced prolonged droughts and poor rainfall distribution, while the dambos have experienced recurrent seasonal floods. This study assessed some of the adaptive strategies exercised by small-scale rural farmers in response to climate change in the Shire Valley. The methodology used in collecting information includes group discussions, household surveys in the area, secondary data, and field observations. The results show that small-scale rural farmers exercise a number of adaptive strategies in response to climate change. These adaptive strategies include: increased use of water resources for small-scale irrigation or wetland farming, mostly using simple delivery techniques; increased management of residual moisture; and increased alternative sources of income such as fishing and crop diversity. It was also observed that government promoted the use of portable motorized pumps for small-scale irrigation in order to mitigate the effects of climate change. However, these external interventions were not fully adopted; instead the farmers preferred local interventions which mostly had indigenous elements.
Although HIV-prevalence and fertility rates in sub-Saharan Africa are among the highest in the world, little is known about how HIV infection affects the fertility preferences of men and women in the region. A quasi-experimental design and in-depth interviews conducted in rural Malawi are employed to examine how and through what pathways learning that one is HIV positive alters a person’s childbearing desires. Among rural Malawians, particularly men, the desire to have more children decreases after receiving a positive HIV-test result. The motivations underlying this effect are greatly influenced by gender: women fear the physical health consequences of HIV-positive pregnancies and childbearing, whereas men see childbearing as futile because they anticipate their own early death and the deaths of their future children. Considerable ambivalence remains, nevertheless, particularly among women who strategize to live normal lives in spite of their infection, but whose definitions of “normal” vary. PMID:21151844
Thomas, Kevin J. A.; Inkpen, Christopher
Using data from Malawi, this study situates the discourse on migration, entrepreneurship, and development within the context of Africa’s social realities. It examines self-employment differences among three groups of migrants and corresponding group differences in agricultural and non-agricultural self-employment. International migrants are found to be more engaged in self-employment than internal-migrants. However, our results suggest that previous findings on the development-related contributions of returning migrants from the West need to be appropriately contextualized. When returnees from the West invest in self-employment, they typically shy away from Africa’s largest economic sector - agriculture. In contrast, levels of self-employment, especially in agricultural self-employment, are highest among returning migrants and immigrants from other African countries, especially from those nearby. We also underscore the gendered dimensions of migrants’ contribution to African development by demonstrating that female migrants are more likely to be self-employed in agriculture than male migrants. Furthermore, as human-capital increases, migrants are more likely to concentrate their self-employment activities in non-agricultural activities and not in the agricultural sector. The study concludes by using these findings to discuss key implications for policy and future research. PMID:25598569
Jagger, Pamela; Perez-Heydrich, Carolina
Interventions to mitigate household air pollution (HAP) from cooking with solid fuels often fail to take into account the role of access to freely available woodfuels in determining fuel choice and willingness to adopt clean cooking technologies, key factors in mitigating the burden of HAP. We use national-scale remote sensing data on land use land cover change, and population representative data from two waves of the Malawi Living Standards Measurement Survey to explore the relationship between land use change and the type of fuel households use, time spent collecting fuel, and expenditures on fuel, hypothesizing that land use dynamics influence household-level choice of primary cooking fuel. We find considerable heterogeneity with respect to regeneration and deforestation/degradation dynamics and evidence of spatial clustering. We find that regeneration of forests and woodlands increases the share of households that collect fuelwood, whereas deforestation and degradation lead households to purchase fuelwood. We also find that a relatively large share of land under woody savannah or degraded forest (versus fully stocked forest) increases fuel collection time. Areas with regeneration happening at broader scale experience increases in fuel expenditures. Our findings have implications for the spatial targeting of interventions designed to mitigate HAP.
Grant, Monica J.; Lloyd, Cynthia B.; Mensch, Barbara S.
The provision of toilets and menstrual supplies has emerged as a promising programmatic strategy to support adolescent girls’ school attendance and performance in less developed countries. We use the first round of the Malawi Schooling and Adolescent Survey (MSAS) to examine the individual- and school-level factors associated with menstruation-related school absenteeism. The MSAS is a school-based longitudinal survey of adolescent students enrolled in coed public primary schools in the southern districts of Machinga and Balaka who were aged 14–16 in 2007. Although one-third of female students report missing at least one day of school at their last menstrual period, our data suggest that menstruation only accounts for a small proportion of all female absenteeism and does not create a gender gap in absenteeism. We find no evidence for school-level variance in menstruation-related absenteeism, suggesting that absenteeism is not sensitive to school environments. Rather, co-residence with a grandmother and spending time on school work at home reduce the odds of absence during the last menstrual period. PMID:25580018
Shacham, Enbal; Thornton, Rebecca; Godlonton, Susan; Murphy, Ryan; Gilliland, Jake
Prevalence of HIV in sub-Saharan African countries persists at alarming rates. There are currently four promoted methods to prevent HIV infection: adherence to antiretroviral therapy, male circumcision, pre-exposure prophylaxis and use of condoms. This study aimed to assess the availability and accessibility of one of the prevention efforts, condoms, in Kawale, Lilongwe, Malawi. A total of 220 potential condom-selling establishments were surveyed in 2012. Data were collected with store owners or staff and locations were geocoded to assess store density. Descriptive analyses were conducted. Of those audited, 96 stores sold condoms, 13 of which distributed free condoms. The stores were most often small shops and located in markets or trading centres. Condoms were most often found at the back of the store in an open space. There were approximately 1.2 stores per ¼ mile; 44% of the businesses in the study region carried condoms. This one method of prevention exhibited multiple barriers in this region: few stores sold condoms, high costs, condom locations within stores and limited availability. The limited accessibility is likely to influence social norms surrounding condom use. Future research should incorporate assessing norms and addressing barriers to uptake of HIV prevention efforts.
Das, Ipsita; Jagger, Pamela; Yeatts, Karin
In sub-Saharan Africa, biomass fuels account for approximately 90% of household energy consumption. Limited evidence exists on the association between different biomass fuels and health outcomes. We report results from a cross-sectional sample of 655 households in Malawi. We calculated odds ratios between hypothesized determinants of household air pollution (HAP) exposure (fuel, stove type, and cooking location) and five categories of health outcomes (cardiopulmonary, respiratory, neurologic, eye health, and burns). Reliance on high- or low-quality firewood or crop residue (vs. charcoal) was associated with significantly higher odds of shortness of breath, difficulty breathing, chest pains, night phlegm, forgetfulness, dizziness, and dry irritated eyes. Use of high-quality firewood was associated with significantly lower odds of persistent phlegm. Cooks in rural areas (vs. urban areas) had significantly higher odds of experiencing shortness of breath, persistent cough, and phlegm, but significantly lower odds of phlegm, forgetfulness, and burns. With deforestation and population pressures increasing reliance on low-quality biomass fuels, prevalence of HAP-related cardiopulmonary and neurologic symptoms will likely increase among cooks. Short- to medium-term strategies are needed to secure access to high-quality biomass fuels given limited potential for scalable transitions to modern energy.
Thomas, Kevin J A; Inkpen, Christopher
Using data from Malawi, this study situates the discourse on migration, entrepreneurship, and development within the context of Africa's social realities. It examines self-employment differences among three groups of migrants and corresponding group differences in agricultural and non-agricultural self-employment. International migrants are found to be more engaged in self-employment than internal-migrants. However, our results suggest that previous findings on the development-related contributions of returning migrants from the West need to be appropriately contextualized. When returnees from the West invest in self-employment, they typically shy away from Africa's largest economic sector - agriculture. In contrast, levels of self-employment, especially in agricultural self-employment, are highest among returning migrants and immigrants from other African countries, especially from those nearby. We also underscore the gendered dimensions of migrants' contribution to African development by demonstrating that female migrants are more likely to be self-employed in agriculture than male migrants. Furthermore, as human-capital increases, migrants are more likely to concentrate their self-employment activities in non-agricultural activities and not in the agricultural sector. The study concludes by using these findings to discuss key implications for policy and future research.
McCartney, Tannis; Scholz, Christopher A.
This paper analyzes throw-depth (T-z) profiles from a high resolution 2D reflection seismic grid in the central basin of Lake Malawi to investigate whether evidence exists: 1) for migration of faulting away from the border fault of the half-graben; and 2) that faults in the hangingwall lengthened over the last 1.3 million years. We use the high-precision age model from a 2005 scientific drilling project in our study area to constrain the ages of our seismic horizons and examine a fault array and two individual faults within the hangingwall of the central basin border fault. We account for climatic and sedimentological controls on stratal growth with a lake-level curve that accompanies the age model. A comparison of our hangingwall T-z profiles with published throw-distance (T-x) profiles for the border fault shows synchronous faulting over the last 1.3 m.y. rather than basinward migration of faulting. Furthermore, we find no evidence for significant propagation of the tips of the hangingwall faults in the last 1.3 m.y. and conclude that the lack of basinward migration of faulting is a consequence of strain localization on faults established at an early stage in basin development.
Mazzuchelli, Juliana; Yang, Fengtang; Kocher, Thomas D.; Martins, Cesar
To better understand the genomic organization and evolution of Sox genes in vertebrates, we cytogenetically mapped Sox2 and Sox14 genes in cichlid fishes and performed comparative analyses of their orthologs in several vertebrate species. The genomic regions neighbouring Sox2 and Sox14 have been conserved during vertebrate diversification. Although cichlids seem to have undergone high rates of genomic rearrangements, Sox2 and Sox14 are linked in the same chromosome in the Etroplinae Etroplus maculatus that represents the sister group of all remaining cichlids. However, this genes are located on different chromosomes in several species of the sister group Pseudocrenilabrinae. Similarly the ancestral synteny of Sox2 and Sox14 has been maintained in several vertebrates, but this synteny has been broken independently in all major groups as a consequence of karyotype rearrangements that took place during the vertebrate evolution. PMID:21691861
Theis, Anya; Salzburger, Walter; Egger, Bernd
Color and pigmentation patterns of animals are often targets of sexual selection because of their role in communication. Although conspicuous male traits are typically implicated with intersexual selection, there are examples where sex-specific displays play a role in an intrasexual context, e.g. when they serve as signals for aggression level and/or status. Here, we focus on the function of a conspicuous male ornament in the most species-rich tribe of cichlid fishes, the haplochromines. A characteristic feature of these ca. 1500 species are so-called egg-spots in form of ovoid markings on the anal fins of males, which are made up of carotenoid based pigment cells. It has long been assumed that these yellow, orange or reddish egg-spots play an important role in the courtship and spawning behavior of these maternal mouth-brooding fishes by mimicking the eggs of a conspecific female. The exact function of egg-spots remains unknown, however, and there are several hypotheses about their mode of action. To uncover the function of this cichlid-specific male ornament, we used female mate choice experiments and a male aggression test in the haplochromine species Astatotilapia burtoni. We manipulated the number and arrangement of egg-spots on the anal fins of males, or removed them entirely, and tested (1) female preference with visual contact only using egg-traps, (2) female preference with free contact using paternity testing with microsatellites and (3) male aggression. We found that females did not prefer males with many egg-spots over males with fewer egg-spots and that females tended to prefer males without egg-spots over males with egg-spots. Importantly, males without egg-spots sired clutches with the same fertilization rate as males with egg-spots. In male aggression trials, however, males with fewer egg-spots received significantly more attacks, suggesting that egg-spots are an important signal in intrasexual communication. PMID:22242184
Izugbara, Chimaraoke O.; Undie, Chi-Chi; Mudege, Netsayi N.; Ezeh, Alex C.
There is limited research addressing the beliefs of adolescents related to Voluntary Counseling and HIV-Testing (VCT). This paper analyzes qualitative data on such beliefs elicited from male youth in Uganda and Malawi. Although study participants understood the mainstream public health rhetoric on VCT, much of their narratives framed going for HIV…
Wellard, Kate; Rafanomezana, Jenny; Nyirenda, Mahara; Okotel, Misaki; Subbey, Vincent
Purpose: Farmer-to-farmer extension offers a potentially low-cost and wide-reach alternative in supporting agricultural innovation. Various approaches are being promoted but information on their impact and sustainability is sparse. This study examines experiences of Self Help Africa and partners in Ghana, Uganda and Malawi. It asks: What is good…
International Planned Parenthood Federation, London (England).
Data relating to population and family planning in eight foreign countries are presented in these situation reports. Countries included are Hong Kong, Malawi, Pakistan, Panama, Philippines, Ryukyu Islands (Okinawa), Sabah, and Sarawak. Information is provided where appropriate and available, under two topics, general background and family planning…
Gombachika, Harry S. H.; Khangamwa, Gift
Purpose: The paper examines the effects of ICT readiness dimensions on ICT acceptance among Technical, Entrepreneurial and Vocational Training (TEVT) students in the University of Malawi, using correlation and regression analyses. Design/methodology/approach: A cross-sectional survey was used with a sample of 125 respondents drawn from a…
Shawa, Lester B.; Mgomezulu, Victor Y.
Utilising critical theory, we explored the causes of the conflict that arose between academic staff on fixed-term renewable contracts and university administrators at Mzuzu University in Malawi in order to draw lessons. We collected data using semi-structured, in-depth interviews and document analysis. Ten university employees were purposively…
Educational reforms, particularly in a contested subject such as Religious Education (RE), have unsettled boundaries principally because actors demand or expect different outcomes of these reforms. In the cases of Scotland and Malawi the present paper examines how different stakeholders have engaged with RE reforms. It thus ascertains whether, if…
Peace Corps, Washington, DC.
This vocabulary guide is an alphabetical listing of English words and their equivalent in Chinyanja. A general list of verbs is included, as well as commonly used expressions for planned conversation. The basic expressions are intended for use in learning the health needs and problems of the Malawi community. (NCR)
Glasson, George E.; Frykholm, Jeffrey A.; Mhango, Ndalapa A.; Phiri, Absalom D.
The purpose of this 2-year study was to investigate Malawian teacher educators' perspectives and dispositions toward teaching about ecological sustainability issues in Malawi, a developing country in sub-Sahara Africa. This study was embedded in a larger theoretical framework of investigating earth systems science through the understanding of…
Kaponda, Chrissie P. N.; Norr, Kathleen F.; Crittenden, Kathleen S.; Norr, James L.; McCreary, Linda L.; Kachingwe, Sitingawawo I.; Mbeba, Mary M.; Jere, Diana L. N.; Dancy, Barbara L.
This study used a quasi-experimental design to evaluate a six-session peer group intervention for HIV prevention among rural adults in Malawi. Two rural districts were randomly assigned to intervention and control conditions. Independent random samples of community adults compared the districts at baseline and at 6 and 18 months postintervention.…
Holland, Dana G.
Changes in research production precipitated by the globalization have generally been theorized as applying across nations and disciplinary projects. This article examines the relation of discipline to research production from the situational vantage point of the developing world, specifically the Southern African country of Malawi, and from the…
Kamwendo, Gregory Hankoni
This research note is a critique of Malawi's new language of instruction (LOI) policy. The new policy stipulates English as the medium of instruction from the first year (Standard 1) of primary school in a country where English is not the dominant language of household communication. The children are now expected to learn (and that includes…
Glasson, George E.; Frykholm, Jeffrey A.; Mhango, Ndalapa A.; Phiri, Absalom D.
The purpose of this 2-year study was to investigate Malawian teacher educators' perspectives and dispositions toward teaching about ecological sustainability issues in Malawi, a developing country in sub-Sahara Africa. This study was embedded in a larger theoretical framework of investigating earth systems science through the understanding of nature-knowledge-culture systems from local, place-based perspectives. Specifically, we were interested in learning more about eco-justice issues that are related to environmental degradation in Malawi and the potential role of inquiry-oriented pedagogies in addressing these issues. In a science methods course, the African educators' views on deforestation and teaching about ecological sustainability were explored within the context of the local environment and culture. Teachers participated in inquiry pedagogies designed to promote the sharing of perspectives related to the connections between culture and ecological degradation. Strategies encouraging dialogue and reflection included role-playing, class discussions, curriculum development activities, teaching experiences with children, and field trips to a nature preserve. Data were analyzed from postcolonial and critical pedagogy of place theoretical perspectives to better understand the hybridization of viewpoints influenced by both Western and indigenous science and the political hegemonies that impact sustainable living in Malawi. Findings suggested that the colonial legacy of Malawi continues to impact the ecological sustainability issue of deforestation. Inquiry-oriented pedagogies and connections to indigenous science were embraced by the Malawian educators as a means to involve children in investigation, decision making, and ownership of critical environmental issues.
McCreary, Linda L.; Kaponda, Chrissie P. N.; Kafulafula, Ursula K.; Ngalande, Rebecca C.; Kumbani, Lily C.; Jere, Diana L. N.; Norr, James L.; Norr, Kathleen F.
This paper reports the process evaluation of a peer group intervention for human immunodeficiency virus (HIV) prevention which had positive outcomes for three target groups in Malawi: rural adults, adolescents and urban hospital workers. The six-session intervention was delivered to small groups of 10-12 participants by 85 trained volunteer peer…
Inoue, Kazuma; Oketch, Moses
Malawi and Ghana are among the numerous Sub-Saharan Africa countries that have in recent years introduced Free Primary Education (FPE) policy as a means to realizing the 2015 Education for All and Millennium Development Goals international targets. The introduction of FPE policy is, however, a huge challenge for any national government that has…
Nampota, Dorothy; Thompson, Jeff; Wikeley, Felicity
Faced with accelerating poverty, the Malawi government has re-awakened its commitment to the development of human capacity and the role of development in this context. This paper explores the relationship between development and science and technology. It goes on to review the country's science and technology needs and how these justify taking…
Lubeski, Nancy J.
This doctoral research work explores the quality of educational experiences and access to opportunities for girls with disabilities in selected rural primary schools in Malawi. The context for this study lies within four of the six goals contained within the World Declaration on Education for All doctrine, a document which lays out a framework for…
Stockman, Lauren J; Fischer, Thea K; Deming, Michael; Ngwira, Bagrey; Bowie, Cameron; Cunliffe, Nigel; Bresee, Joseph; Quick, Robert E
A national household survey was conducted in Malawi to determine awareness and use of a socially marketed water treatment product. In all, 64% of mothers were aware of the product, and 7% were using it. Both poor and rural mothers had lower awareness and use rates. Targeting promotion to rural populations could enhance program effectiveness.
White Lake is in Muskegon County along the eastern shore of Lake Michigan. It was named an Area of Concern on the Great Lakes under the Great Lakes Water Quality Agreement of 1987 and delisted in 2014.
Chunga, Richard M; Ensink, Jeroen H J; Jenkins, Marion W; Brown, Joe
This paper presents the results of a mixed-methods study examining adaptation strategies that property owners in low-income, rapidly urbanizing areas in Malawi adopt to address the limitations of pit latrines, the most common method of disposing human excreta. A particular challenge is lack of space for constructing new latrines as population density increases: traditional practice has been to cap full pits and simply move to a new site, but increasing demands on space require new approaches to extend the service life of latrines. In this context, we collected data on sanitation technology choices from January to September 2013 through 48 in-depth interviews and a stated preference survey targeting 1,300 property owners from 27 low-income urban areas. Results showed that property owners with concern about space for replacing pit latrines were 1.8 times more likely to select pit emptying service over the construction of new pit latrines with a slab floor (p = 0.02) but there was no significant association between concern about space for replacing pit latrines and intention to adopt locally promoted, novel sanitation technology known as ecological sanitation (ecosan). Property owners preferred to adapt existing, known technology by constructing replacement pit latrines on old pit latrine locations, reducing the frequency of replacing pit latrines, or via emptying pit latrines when full. This study highlights potential challenges to adoption of wholly new sanitation technologies, even when they present clear advantages to end users. To scale, alternative sanitation technologies for rapidly urbanising cities should offer clear advantages, be affordable, be easy to use when shared among multiple households, and their design should be informed by existing adaptation strategies and local knowledge.
Msonda, K. W. M.; Masamba, W. R. L.; Fabiano, E.
A study was carried out to determine fluoride concentration in groundwaters of Nathenje area situated in Lilongwe District in the central region of Malawi. Water samples were collected from 176 boreholes and shallow wells during different months in 2001 and 2002. Samples were then analysed for fluoride by using a fluoride electrode and an ion selective meter. The results showed that fluoride concentrations for the rainy season varied from <0.5 to 6.98 ± 0.01 mg/l with 52.9% of the boreholes above the World Health Organisation (WHO) maximum permissible limit of 1.5 mg/l. Fluoride concentrations for dry season ranged from <0.5 to 7.02 ± 0.02 mg/l with 50.8% of boreholes above 1.5 mg/l. Fluoride concentrations for the two seasons were significantly different from each other ( p < 0.05). Fluoride data was used to produce a fluoride distribution map. From the map, it was observed that fluoride concentrations in this area followed a pattern. The central part of Nathenje had high fluoride concentration of between 2 and 7.02 mg/l and these high fluoride values seemed to extend eastwards beyond the boundary of the study area. However, the southern and western parts had <1 mg/l of fluoride. The high groundwater fluoride values seem to be associated with the weathered basement complex containing biotite that is a probable source of fluoride. The other suspected sources of fluoride in Nathenje groundwater could be due to the dissolution of hornblende, fluorite and amphibole, which are reported to occur in rocks and soils in this area. There was evidence of dental fluorosis in areas where the fluoride concentration was high.
Chunga, Richard M.; Ensink, Jeroen H. J.; Jenkins, Marion W.; Brown, Joe
This paper presents the results of a mixed-methods study examining adaptation strategies that property owners in low-income, rapidly urbanizing areas in Malawi adopt to address the limitations of pit latrines, the most common method of disposing human excreta. A particular challenge is lack of space for constructing new latrines as population density increases: traditional practice has been to cap full pits and simply move to a new site, but increasing demands on space require new approaches to extend the service life of latrines. In this context, we collected data on sanitation technology choices from January to September 2013 through 48 in-depth interviews and a stated preference survey targeting 1,300 property owners from 27 low-income urban areas. Results showed that property owners with concern about space for replacing pit latrines were 1.8 times more likely to select pit emptying service over the construction of new pit latrines with a slab floor (p = 0.02) but there was no significant association between concern about space for replacing pit latrines and intention to adopt locally promoted, novel sanitation technology known as ecological sanitation (ecosan). Property owners preferred to adapt existing, known technology by constructing replacement pit latrines on old pit latrine locations, reducing the frequency of replacing pit latrines, or via emptying pit latrines when full. This study highlights potential challenges to adoption of wholly new sanitation technologies, even when they present clear advantages to end users. To scale, alternative sanitation technologies for rapidly urbanising cities should offer clear advantages, be affordable, be easy to use when shared among multiple households, and their design should be informed by existing adaptation strategies and local knowledge. PMID:27532871
Zere, Eyob; Moeti, Matshidiso; Kirigia, Joses; Mwase, Takondwa; Kataika, Edward
Background Growing scientific evidence points to the pervasiveness of inequities in health and health care and the persistence of the inverse care law, that is the availability of good quality healthcare seems to be inversely related to the need for it in developing countries. Achievement of the Millennium Development Goals is likely to be compromised if inequities in health/healthcare are not properly addressed. Objective This study attempts to assess trends in inequities in selected indicators of health status and health service utilization in Malawi using data from the Demographic and Health Surveys of 1992, 2000 and 2004. Methods Data from Demographic and Health Surveys of 1992, 2000 and 2004 are analysed for inequities in health/healthcare using quintile ratios and concentration curves/indices. Results Overall, the findings indicate that in most of the selected indicators there are pro-rich inequities and that they have been widening during the period under consideration. Furthermore, vertical inequities are observed in the use of interventions (treatment of diarrhoea, ARI among under-five children), in that the non-poor who experience less burden from these diseases receive more of the treatment/interventions, whereas the poor who have a greater proportion of the disease burden use less of the interventions. It is also observed that the publicly provided services for some of the selected interventions (e.g. child delivery) benefit the non-poor more than the poor. Conclusion The widening trend in inequities, in particular healthcare utilization for proven cost-effective interventions is likely to jeopardize the achievement of the Millennium Development Goals and other national and regional targets. To counteract the inequities it is recommended that coverage in poor communities be increased through appropriate targeting mechanisms and effective service delivery strategies. There is also a need for studies to identify which service delivery mechanisms are
Bachan, Lauren K.
BACKGROUND While there is a rich literature on the practice of child fostering in sub-Saharan Africa, little is known about how fostering impacts receiving households, as few studies consider household conditions both before and after fostering. Despite the fact that circumstances surrounding fostering vary, the literature’s key distinction of fostering is often drawn along the simple line of whether or not a household is fostering a child. This paper argues that anticipation of fostering responsibilities, in particular, is a useful dimension to distinguish fostering experiences for receiving households. OBJECTIVE This paper examines the relationship between receiving a foster child and subsequent changes in household wealth. Particular emphasis is placed on how these changes are conditioned by differing levels of anticipation of the fostering event. METHODS This study uses data from Tsogolo la Thanzi (TLT), a longitudinal survey in Balaka, Malawi. Using data from 1754 TLT respondents, fixed effects pooled time-series models are estimated to assess whether and how receiving a foster child changes household wealth. RESULTS This paper demonstrates the heterogeneity of fostering experiences for receiving households. The results show that households that anticipate fostering responsibilities experience a greater increase in household wealth than both households that do not foster and those that are surprised by fostering. CONCLUSION Households that anticipate fostering responsibilities exhibit the greatest increase in household wealth. While fostering households that do not anticipate fostering responsibilities may not experience these gains, there is no evidence to indicate that such households are negatively impacted relative to households that do not foster. This finding suggests that additional childcare responsibilities may not be as detrimental to African households as some researchers have feared. PMID:25419172
Nagl, S; Tichy, H; Mayer, W E; Takezaki, N; Takahata, N; Klein, J
According to a widely held view, the more than 300 species of haplochromine cichlid fishes in Lake Victoria (LV), East Africa, originated from a single founder species in less than 12,000 years. This view, however, does not follow from the published geological and molecular evidence. The former does indeed suggest that the LV basin dried out less than 15,000 years ago, but it does not provide any information about the species that re-colonized the new lake or that remained in the rivers draining the area. The molecular evidence is inconclusive with respect to the origin of the LV haplochromines because cichlids from critical regions around LV were not adequately sampled; and as far as the age of the LV haplochromines is concerned, it in fact led to an estimate of 250,000-750,000 years old. In the present study, mitochondrial DNA (control region) variation was determined by heteroduplex and sequencing analyses of more than 670 specimens collected at widely distributed East African riverine and lacustrine localities. The analyses revealed the existence of seven haplogroups (I-VII) distinguishable by characteristic substitutions. All endemic LV samples tested fell into one of these haplogroups (V) which, however, was also found to be present at various other localities, both riverine and lacustrine, outside LV. Within this haplogroup, four subgroups (VA through VD) could be distinguished, two of which (VB and VC) were represented in LV and at other localities. The great majority of the LV haplochromine species could be classified as belonging to the VC subgroup, which was found only in LV and in the rivers draining into it. Hence, while the endemic haplochromine species of LV could not have originated from a single founding population, the lake does harbour a large species flock which probably arose in situ.
Cooperative breeders serve as a model to study the evolution of cooperation, where costs and benefits of helping are typically scrutinized at the level of group membership. However, cooperation is often observed in multi-level social organizations involving interactions among individuals at various levels. Here, we argue that a full understanding of the adaptive value of cooperation and the evolution of complex social organization requires identifying the effect of different levels of social organization on direct and indirect fitness components. Our long-term field data show that in the cooperatively breeding, colonial cichlid fish Neolamprologus pulcher, both large group size and high colony density significantly raised group persistence. Neither group size nor density affected survival at the individual level, but they had interactive effects on reproductive output; large group size raised productivity when local population density was low, whereas in contrast, small groups were more productive at high densities. Fitness estimates of individually marked fish revealed indirect fitness benefits associated with staying in large groups. Inclusive fitness, however, was not significantly affected by group size, because the direct fitness component was not increased in larger groups. Together, our findings highlight that the reproductive output of groups may be affected in opposite directions by different levels of sociality, and that complex forms of sociality and costly cooperation may evolve in the absence of large indirect fitness benefits and the influence of kin selection. PMID:26582022
McCafferty, S. Shawn; Martin, Andrew; Bermingham, Eldredge
It is well appreciated that historical and ecological processes are important determinates of freshwater biogeographic assemblages. Phylogeography can potentially lend important insights into the relative contribution of historical processes in biogeography. However, the extent that phylogeography reflects historical patterns of drainage connection may depend in large part on the dispersal capability of the species. Here, we test the hypothesis that due to their relatively greater dispersal capabilities, the neotropical cichlid species Andinoacara coeruleopunctatus will display a phylogeographic pattern that differs from previously described biogeographic assemblages in this important region. Based on an analysis of 318 individuals using mtDNA ATPase 6/8 sequence and restriction fragment length polymorphism data, we found eight distinct clades that are closely associated with biogeographic patterns. The branching patterns among the clades and a Bayesian clock analysis suggest a relatively rapid colonization and diversification among drainages in the emergent Isthmus of Panama followed by the coalescing of some drainages due to historical connections. We also present evidence for extensive cross-cordillera sharing of clades in central Panama and the Canal region. Our results suggest that contemporary phylogeographic patterns and diversification in Lower Central American fishes reflect an interaction of historical drainage connections, dispersal, and demographic processes. PMID:23008800
Schneider, Carlos Henrique; Gross, Maria Claudia; Terencio, Maria Leandra; de Tavares, Édika Sabrina Girão Mitozo; Martins, Cesar; Feldberg, Eliana
Fish of the family Cichlidae are recognized as an excellent model for evolutionary studies because of their morphological and behavioral adaptations to a wide diversity of explored ecological niches. In addition, the family has a dynamic genome with variable structure, composition and karyotype organization. Microsatellites represent the most dynamic genomic component and a better understanding of their organization may help clarify the role of repetitive DNA elements in the mechanisms of chromosomal evolution. Thus, in this study, microsatellite sequences were mapped in the chromosomes of Cichla monoculus Agassiz, 1831, Pterophyllum scalare Schultze, 1823, and Symphysodon discus Heckel, 1840. Four microsatellites demonstrated positive results in the genome of Cichla monoculus and Symphysodon discus, and five demonstrated positive results in the genome of Pterophyllum scalare. In most cases, the microsatellite was dispersed in the chromosome with conspicuous markings in the centromeric or telomeric regions, which suggests that sequences contribute to chromosome structure and may have played a role in the evolution of this fish family. The comparative genome mapping data presented here provide novel information on the structure and organization of the repetitive DNA region of the cichlid genome and contribute to a better understanding of this fish family's genome.
Oliveira, Tânia F.; Ros, Albert F. H.; Oliveira, Rui F.
In many territorial species androgen hormones are known to increase in response to territorial intrusions as a way to adjust the expression of androgen-dependent behaviour to social challenges. The dear enemy effect has also been described in territorial species and posits that resident individuals show a more aggressive response to intrusions by strangers than by other territorial neighbours. Therefore, we hypothesized that the dear enemy effect may also modulate the androgen response to a territorial intrusion. Here we tested this hypothesis in male cichlid fish (Mozambique tilapia, Oreochromis mossambicus) using a paradigm of four repeated territorial intrusions, either by the same neighbour or by four different unfamiliar intruders. Neighbour intruders elicited lower aggression and a weaker androgen response than strangers on the first intrusion of the experiment. With repeated intrusions, the agonistic behaviour of the resident males against familiar intruders was similar to that displayed towards strangers. By the fourth intrusion the androgen response was significantly reduced and there was no longer a difference between the responses to the two types of intruders. These results suggest that the dear enemy effect modulates the androgen response to territorial intrusions and that repeated intrusions lead to a habituation of the androgen response. PMID:26379045
Kohda, Masanori; Heg, Dik; Makino, Yoshimi; Takeyama, Tomohiro; Shibata, Jun-ya; Watanabe, Katsutoshi; Munehara, Hiroyuki; Hori, Michio; Awata, Satoshi
Theories suggest that, in cooperatively breeding species, female control over paternity and reproductive output may affect male reproductive skew and group stability. Female paternity control may come about through cryptic female choice or female reproductive behaviour, but experimental studies are scarce. Here, we show a new form of female paternity control in a cooperatively polyandrous cichlid fish (Julidochromis transcriptus), in which females prefer wedge-shaped nesting sites. Wedge-shaped sites allowed females to manipulate the siring success of the group member males by spawning the clutch at the spot where the large males were just able to enter and fertilize the outer part of the clutch. Small males fertilized the inner part of the clutch, protected from the large aggressive males, leading to low male reproductive skew. Small males provided more brood care than large males. Multiple paternity induced both males to provide brood care and reduced female brood care accordingly. This is, to our knowledge, the first documented case in a species with external fertilization showing female mating behaviour leading to multiple male paternity and increased male brood care as a result.
Kidd, Michael R; O'Connell, Lauren A; Kidd, Celeste E; Chen, Christine W; Fontenot, Miles R; Williams, Sidney J; Hofmann, Hans A
Mate choice is fundamental to sexual selection, yet little is known about underlying physiological mechanisms that influence female mating decisions. We investigated the endocrine underpinnings of female mate choice in the African cichlid Astatotilapia burtoni, a non-seasonal breeder. In addition to profiling behavioral and hormonal changes across the female reproductive cycle, we tested two hypotheses regarding possible factors influencing female mate choice. We first asked whether female mate choice is influenced by male visual and/or chemical cues. A. burtoni females were housed for one full reproductive cycle in the center of a dichotomous choice apparatus with a large (attractive) or small (unattractive) conspecific male on either side. Females associated mostly with small, less attractive males, but on the day of spawning reversed their preference to large, attractive males, with whom they mated almost exclusively, although this choice depended on the relative amount of androgens released into the water by small males. We next asked whether male behavior or androgen levels change in relation to the stimulus females' reproductive state. We found that stimulus male aggression decreased and reproductive displays increased as the day of spawning approached. Moreover male testosterone levels changed throughout the females' reproductive cycle, with larger males releasing more testosterone into the water than small males. Our data suggest that female association in a dichotomous choice assay is only indicative of the actual mate choice on the day of spawning. Furthermore, we show that male behavior and hormone levels are dependent on the reproductive state of conspecific females.
Oliveira, R F; Almada, V C; Canario, A V
The relationship between urinary concentrations (free + sulfates + glucuronides) of the steroids testosterone (T), 11-ketotestosterone (11KT), 17 alpha, 20 beta-dihydroxy-4-pregnen-3-one (17, 20 beta-P) and 17 alpha, 20 alpha-dihydroxy-4-pregnen-3-one (17, 20 alpha-P), and the social behavior of males of the cichlid fish Oreochromis mossambicus was investigated. After 8 days of isolation none of the steroids were good predictors of social dominance developed after subsequent formation of all-male groups. One day after group formation dominance indexes were good predictors of the urine concentrations of all sex steroids. Dominance indexes and androgen concentrations measured after all-male group formation were positively correlated with territoriality, courtship rate, and nest size. Similar relationships were found for progestins with the exception that they were not correlated with courtship rate. All-male group formation was also accompanied by an increase in urinary sex steroid concentrations in fish that became territorial and a decrease in non-territorial fish with the exception of T, which increased in both groups. Addition of ovulating females caused steroid concentrations to return to levels near isolation, except for 17, 20 alpha-P in territorials, which underwent a large increase. Thus, social interactions may have an important modulatory effect on sex steroid concentrations in O. mossambicus.
Mesterton-Gibbons, Mike; Dai, Yao
We determine size, shape and location for a territory that is optimal in the sense of minimizing defense costs, when a given proportion of the boundary is landmarked and its primary benefit in terms of fitness is greater ease of detecting intruders across it. Increasing the landmarked proportion of boundary causes the optimal configuration to be smaller and more elongated, and to be located with its center further from the nest, so that the nest is closer to the landmarked boundary. These predictions accord with observations in a recent study of the convict cichlid Amatitlania siquia. Our results thus confirm the consistency of the observed behavior with the hypothesis that A. siquia designs its territory to make intruders easier to spot. Our results also lead us to conjecture that moving the landmark proportionately closer to or further away from the nest would have yielded essentially the same outcome in this study, because the optimal configuration depends only on the angle subtended by the landmark at the nest and hence only on the length of the landmark relative to its distance from the nest, as opposed to its absolute value.
Wirtz Ocana, Sabine; Meidl, Patrick; Bonfils, Danielle; Taborsky, Michael
Behavioural variation among conspecifics is typically contingent on individual state or environmental conditions. Sex-specific genetic polymorphisms are enigmatic because they lack conditionality, and genes causing adaptive trait variation in one sex may reduce Darwinian fitness in the other. One way to avoid such genetic antagonism is to control sex-specific traits by inheritance via sex chromosomes. Here, controlled laboratory crossings suggest that in snail-brooding cichlid fish a single locus, two-allele polymorphism located on a sex-linked chromosome of heterogametic males generates an extreme reproductive dimorphism. Both natural and sexual selection are responsible for exceptionally large body size of bourgeois males, creating a niche for a miniature male phenotype to evolve. This extreme intrasexual dimorphism results from selection on opposite size thresholds caused by a single ecological factor, empty snail shells used as breeding substrate. Paternity analyses reveal that in the field parasitic dwarf males sire the majority of offspring in direct sperm competition with large nest owners exceeding their size more than 40 times. Apparently, use of empty snail shells as breeding substrate and single locus sex-linked inheritance of growth are the major ecological and genetic mechanisms responsible for the extreme intrasexual diversity observed in Lamprologus callipterus.
Kohda, Masanori; Heg, Dik; Makino, Yoshimi; Takeyama, Tomohiro; Shibata, Jun-ya; Watanabe, Katsutoshi; Munehara, Hiroyuki; Hori, Michio; Awata, Satoshi
Theories suggest that, in cooperatively breeding species, female control over paternity and reproductive output may affect male reproductive skew and group stability. Female paternity control may come about through cryptic female choice or female reproductive behaviour, but experimental studies are scarce. Here, we show a new form of female paternity control in a cooperatively polyandrous cichlid fish (Julidochromis transcriptus), in which females prefer wedge-shaped nesting sites. Wedge-shaped sites allowed females to manipulate the siring success of the group member males by spawning the clutch at the spot where the large males were just able to enter and fertilize the outer part of the clutch. Small males fertilized the inner part of the clutch, protected from the large aggressive males, leading to low male reproductive skew. Small males provided more brood care than large males. Multiple paternity induced both males to provide brood care and reduced female brood care accordingly. This is, to our knowledge, the first documented case in a species with external fertilization showing female mating behaviour leading to multiple male paternity and increased male brood care as a result. PMID:19726479
Bose, Aneesh P. H.; O'Connor, Constance M.; Balshine, Sigal
Infanticide and offspring cannibalism are taxonomically widespread phenomena. In some group-living species, a new dominant individual taking over a group can benefit from infanticide if doing so induces potential mates to become reproductively available sooner. Despite widespread observations of infanticide (i.e. egg cannibalism) among fishes, no study has investigated whether egg cannibalism occurs in fishes as a result of group takeovers, or how this type of cannibalism might be adaptive. Using the cooperatively breeding cichlid, Neolamprologus pulcher, we tested whether new unrelated males entering the dominant position in a social group were more likely to cannibalize eggs, and whether such cannibalism would shorten the interval until the female's next spawning. Females spawned again sooner if their broods were removed than if they were cared for. Egg cannibalism occurred frequently after a group takeover event, and was rarer if the original male remained with the group. While dominant breeder females were initially highly aggressive towards newcomer males that took over the group, the degree of resistance depended on relative body size differences between the new pair and, ultimately, female aggression did not prevent egg cannibalism. Egg cannibalism, however, did not shorten the duration until subsequent spawning, or increase fecundity during subsequent breeding in our laboratory setting. Our results show that infanticide as mediated through group takeovers is a taxonomically widespread behaviour.
Roche, Dominique G; Leung, Brian; Franco, Edgar F Mendoza; Torchin, Mark E
Empirical studies suggest that most exotic species have fewer parasite species in their introduced range relative to their native range. However, it is less clear how, ecologically, the loss of parasite species translates into a measurable advantage for invaders relative to native species in the new community. We compared parasitism at three levels (species richness, abundance and impact) for a pair of native and introduced cichlid fishes which compete for resources in the Panama Canal watershed. The introduced Nile tilapia, Oreochromis niloticus, was infected by a single parasite species from its native range, but shared eight native parasite species with the native Vieja maculicauda. Despite acquiring new parasites in its introduced range, O. niloticus had both lower parasite species richness and lower parasite abundance compared with its native competitor. There was also a significant negative association between parasite load (abundance per individual fish) and host condition for the native fish, but no such association for the invader. The effects of parasites on the native fish varied across sites and types of parasites, suggesting that release from parasites may benefit the invader, but that the magnitude of release may depend upon interactions between the host, parasites and the environment.
Wirtz Ocana, Sabine; Meidl, Patrick; Bonfils, Danielle; Taborsky, Michael
Behavioural variation among conspecifics is typically contingent on individual state or environmental conditions. Sex-specific genetic polymorphisms are enigmatic because they lack conditionality, and genes causing adaptive trait variation in one sex may reduce Darwinian fitness in the other. One way to avoid such genetic antagonism is to control sex-specific traits by inheritance via sex chromosomes. Here, controlled laboratory crossings suggest that in snail-brooding cichlid fish a single locus, two-allele polymorphism located on a sex-linked chromosome of heterogametic males generates an extreme reproductive dimorphism. Both natural and sexual selection are responsible for exceptionally large body size of bourgeois males, creating a niche for a miniature male phenotype to evolve. This extreme intrasexual dimorphism results from selection on opposite size thresholds caused by a single ecological factor, empty snail shells used as breeding substrate. Paternity analyses reveal that in the field parasitic dwarf males sire the majority of offspring in direct sperm competition with large nest owners exceeding their size more than 40 times. Apparently, use of empty snail shells as breeding substrate and single locus sex-linked inheritance of growth are the major ecological and genetic mechanisms responsible for the extreme intrasexual diversity observed in Lamprologus callipterus. PMID:25232141
Cahn, Matthew D; Brown, Alexandria C; Clotfelter, Ethan D
Vertebrate pigmentation is known to be influenced by oxidative stress, but few studies have tested the hypothesis that structural coloration can be similarly affected. We tested whether fish iridophores, which produce structural color using guanine stacks, might be affected by the prooxidant-antioxidant balance of the animal. Specifically, we hypothesized that convict cichlids (Amatitlania nigrofasciata) metabolize guanine present in iridophores to uric acid, an antioxidant, in response to oxidative damage. We used Hunter's contrast gloss and high performance liquid chromatography to determine whether dietary guanine supplementation allows fish to maintain their structural coloration despite oxidative stress induced via ultraviolet-B (UV-B) radiation. We found that dietary guanine was associated with greater skin gloss, and that exposure to UV-B light reduced glossiness. UV-B exposure did not increase oxidative damage (acrolein) or total antioxidant capacity in the skin or liver. Our experiment did not detect effects of dietary guanine or UV-B light on uric acid, but uric acid was positively related to antioxidant capacity. Our results support the hypothesis that structural color in fish may be altered by environmental stressors such as exposure to UV light, and highlight the need for future studies to consider the role of iridophores in condition-dependent visual signaling.
Hermann, Caroline M.; Brudermann, Verena; Zimmermann, Holger; Vollmann, Johann; Sefc, Kristina M.
Female mate preferences for male traits and resource characteristics affect trait evolution and diversification. Here, we test the effects of male body traits and territory characteristics on within-population female preferences and on population–assortative mating in the cichlid Tropheus moorii. Within-population preferences of females were independent of male body size, coloration and territory size but were strongly dependent on territory quality and co-varied with male courtship activity. Courtship activity of individual males was contingent on the quality of their assigned territory, and therefore, courtship may not only indicate intrinsic male quality. On the basis of these results we suggest that female preferences for high-quality territories reinforce the outcome of malemale competition and ensure male mating success. Mating preferences of females for males of their own color variant (ascertained in a previous experiment) were not overturned when males of another color variant were presented in a superior territory, indicating that within- and between-population mate preferences of females depend on different cues. PMID:25983339
Schneider, Carlos Henrique; Gross, Maria Claudia; Terencio, Maria Leandra; de Tavares, Édika Sabrina Girão Mitozo; Martins, Cesar; Feldberg, Eliana
Abstract Fish of the family Cichlidae are recognized as an excellent model for evolutionary studies because of their morphological and behavioral adaptations to a wide diversity of explored ecological niches. In addition, the family has a dynamic genome with variable structure, composition and karyotype organization. Microsatellites represent the most dynamic genomic component and a better understanding of their organization may help clarify the role of repetitive DNA elements in the mechanisms of chromosomal evolution. Thus, in this study, microsatellite sequences were mapped in the chromosomes of Cichla monoculus Agassiz, 1831, Pterophyllum scalare Schultze, 1823, and Symphysodon discus Heckel, 1840. Four microsatellites demonstrated positive results in the genome of Cichla monoculus and Symphysodon discus, and five demonstrated positive results in the genome of Pterophyllum scalare. In most cases, the microsatellite was dispersed in the chromosome with conspicuous markings in the centromeric or telomeric regions, which suggests that sequences contribute to chromosome structure and may have played a role in the evolution of this fish family. The comparative genome mapping data presented here provide novel information on the structure and organization of the repetitive DNA region of the cichlid genome and contribute to a better understanding of this fish family’s genome. PMID:26753076
Chong, Kenny; Joshi, Shashikant; Jin, Lam Toong; Shu-Chien, Alexander Chong
The discus fish (Symphysodon aequifasciata) is a cichlid demonstrating advanced mode of parental care towards fry. Both male and female fish utilized epidermal mucus secreted from specialized epidermal cells to feed developing fry. We utilized proteomics to compare protein profile from parental and nonparental fish. Gel analysis revealed a total of 35 spots that were up-regulated in parental mucus. In tandem, another 18 spots were uniquely expressed in parental mucus. MS analysis of these spots identified proteins such as fructose biphosphate aldolase, nucleoside diphosphate kinase, and heat shock proteins, which are essential to support energy provision, cell repair and proliferation, stress mediation, and defense mechanism in parental fish during parental-care period. Concurrently, the detection of several antioxidant-related proteins such as thioredoxin peroxidase and hemopexin suggests a need to overcome oxidative stress during hypermucosal production in parental-care behavior. A C-type lectin was also found to be uniquely expressed in parental mucus and could have important role in providing antimicrobial defense to both parental fish and fry. In summary, our study shows that discus mucus proteome undergoes changes in protein expression during parental-care period.
Dijkstra, Peter D; Maguire, Sean M; Harris, Rayna M; Rodriguez, Agosto A; DeAngelis, Ross S; Flores, Stephanie A; Hofmann, Hans A
The melanocortin system is a neuroendocrine system that regulates a range of physiological and behavioural processes. We examined the extent to which the melanocortin system simultaneously regulates colour and behaviour in the cichlid fish Astatotilapia burtoni We found that yellow males are more aggressive than blue males, in line with previous studies. We then found that exogenous α-melanocyte-stimulating hormone (α-MSH) increases yellowness of the body and dispersal of xanthophore pigments in both morphs. However, α-MSH had a morph-specific effect on aggression, with only blue males showing an increase in the rate of aggression. Exogenous agouti signalling peptide (ASIP), a melanocortin antagonist, did not affect coloration but reduced the rate of aggression in both colour morphs. Blue males had higher cortisol levels than yellow males. Neural gene expression of melanocortin receptors (mcr) and ligands was not differentially regulated between colour morphs. In the skin, however, mc1r and pro-opiomelanocortin (pomc) β were upregulated in blue males, while asip 1 was upregulated in yellow males. The effects of α-MSH on behaviour and body coloration, combined with morph-specific regulation of the stress response and the melanocortin system, suggest that the melanocortin system contributes to the polymorphism in behaviour and coloration in A. burtoni.
Jungwirth, Arne; Taborsky, Michael
Cooperative breeders serve as a model to study the evolution of cooperation, where costs and benefits of helping are typically scrutinized at the level of group membership. However, cooperation is often observed in multi-level social organizations involving interactions among individuals at various levels. Here, we argue that a full understanding of the adaptive value of cooperation and the evolution of complex social organization requires identifying the effect of different levels of social organization on direct and indirect fitness components. Our long-term field data show that in the cooperatively breeding, colonial cichlid fish Neolamprologus pulcher, both large group size and high colony density significantly raised group persistence. Neither group size nor density affected survival at the individual level, but they had interactive effects on reproductive output; large group size raised productivity when local population density was low, whereas in contrast, small groups were more productive at high densities. Fitness estimates of individually marked fish revealed indirect fitness benefits associated with staying in large groups. Inclusive fitness, however, was not significantly affected by group size, because the direct fitness component was not increased in larger groups. Together, our findings highlight that the reproductive output of groups may be affected in opposite directions by different levels of sociality, and that complex forms of sociality and costly cooperation may evolve in the absence of large indirect fitness benefits and the influence of kin selection.
Fischer, Stefan; Bessert-Nettelbeck, Mathilde; Kotrschal, Alexander; Taborsky, Barbara
Social animals can greatly benefit from well-developed social skills. Because the frequency and diversity of social interactions often increase with the size of social groups, the benefits of advanced social skills can be expected to increase with group size. Variation in social skills often arises during ontogeny, depending on early social experience. Whether variation of social-group sizes affects development of social skills and related changes in brain structures remains unexplored. We investigated whether, in a cooperatively breeding cichlid, early group size (1) shapes social behavior and social skills and (2) induces lasting plastic changes in gross brain structures and (3) whether the development of social skills is confined to a sensitive ontogenetic period. Rearing-group size and the time juveniles spent in these groups interactively influenced the development of social skills and the relative sizes of four main brain regions. We did not detect a sensitive developmental period for the shaping of social behavior within the 2-month experience phase. Instead, our results suggest continuous plastic behavioral changes over time. We discuss how developmental effects on social behavior and brain architecture may adaptively tune phenotypes to their current or future environments.
Hopperdietzel, C; Hirschberg, R M; Hünigen, H; Wolter, J; Richardson, K; Plendl, J
The primary objectives of this study were to document the macroscopic and histological structure of the alimentary tract (AT) of the convict cichlid Amatitlania nigrofasciata, because there are no data available for this omnivorous freshwater fish of the family Cichlidae. The morphology of the AT of A. nigrofasciata resembles that of related species. While having morphological criteria of the AT typical of most omnivorous fishes, such as a blind sac stomach and medium length intestine, A. nigrofasciata also has some structural peculiarities: the oesophagus is lined by a uniform stratified squamous epithelial layer with interspersed goblet cells along its entire length. Additionally, it has well-developed layers of the tunica muscularis including muscle fibre bundles that ascend into its mucosal folds. Occasionally, taste buds are present. In the transitional area between oesophagus and stomach, a prominent torus-like closure device is present. The mucosa of the stomach cannot be divided into different regions according to mucosal and morphological properties. The simple pattern of intestinal loops of A. nigrofasciata has few variations, irrespective of sex, mass and length of the individual fish. The first segment of the intestine is characterized by the largest mucososerosal ratio and the most complex mucosal surface architecture. A distinction of midgut and hindgut was not possible in A. nigrofasciata due to lack of defining structural components as described for other fish species.
Marques, A C P B; Franco, A C S; Salgueiro, F; García-Berthou, E; Santos, L N
This study used the hypervariable domain of the mitochondrial DNA (mtDNA) control region (CR) to assess the genetic divergence among native and invasive populations of Cichla kelberi, which is considered the first peacock cichlid introduced and established throughout Brazil and is among the most invasive populations of this genus worldwide. The maximum likelihood tree based on 53 CR sequences with strong bootstrap support revealed that C. kelberi forms a monophyletic clade, confirming that all 30 C. kelberi studied belong to this morphotype. Additionally, the haplotype analysis of the C. kelberi sequences from 11 sampling sites revealed that invasive populations are much less diverse than native ones and largely dominated by a single haplotype that prevailed in reservoirs at the Paraíba do Sul River basin. Two haplotypes were recorded exclusively in an invasive population at Porto Rico, southern Brazil, and one private haplotype was detected in two reservoirs from Paraíba do Sul (Pereira Passos and Paracambi), suggesting more than one introduction event and that native populations should be better evaluated to encompass the entire genetic diversity of native C. kelberi. The possible route and pathways of C. kelberi introduction are also briefly discussed.
Anken, Ralf H.; Baur, Ulrich; Hilbig, Reinhard
It has been shown earlier that hypergravity slows down inner ear otolith growth in developing fish as an adaptation towards increased environmental gravity. Suggesting that otolith growth is regulated by the central nervous system, thus adjusting otolithic weight to produce a test mass, applying functional weightlessness should yield an opposite effect, i.e. larger than normal otoliths. Therefore, larval siblings of cichlid fish ( Oreochromis mossambicus) were housed for 7 days in a submersed, two-dimensional clinostat, which provided a residual gravity of approximately 0.007g. After the experiment, otoliths were dissected and their size (area grown during the experiment) was determined. Maintenance in the clinostat resulted in significantly larger utricular otoliths (lapilli, involved in graviperception). There were no statistical significant differences regarding saccular otoliths obtained (sagittae, involved in transmitting linear acceleration and, especially, in the hearing process). These results indicated, that the animals had in fact received functional weightlessness. In line and contrasting results on the otoliths of other teleost species kept at actual microgravity (spaceflight) or within rotating wall vessels are discussed.
Desjardins, Julie K.; Hofmann, Hans A.; Fernald, Russell D.
Social interactions require knowledge of the environment and status of others, which can be acquired indirectly by observing the behavior of others. When being observed, animals can also alter their signals based on who is watching. Here we observed how male cichlid fish (Astatotilapia burtoni) behave when being watched in two different contexts. In the first, we show that aggressive and courtship behaviors displayed by subordinate males depends critically on whether dominant males can see them, and in the second, we manipulated who was watching aggressive interactions and showed that dominant males will change their behavior depending on audience composition. In both cases, when a more dominant individual is out of view and the audience consists of more subordinate individuals, those males signal key social information to females by displaying courtship and dominant behaviors. In contrast, when a dominant male is present, males cease both aggression and courtship. These data suggest that males are keenly aware of their social environment and modulate their aggressive and courtship behaviors strategically for reproductive and social advantage. PMID:22807996
Hotta, Takashi; Takeyama, Tomohiro; Jordan, Lyndon Alexander; Kohda, Masanori
Animal contests are costly and tend to escalate when rivals have similar competitive abilities. Individuals that remember dominance relationships with rivals may avoid repeated agonistic interactions and hence avoid the costs of repeated escalation of contests. However, it can be difficult to experimentally disentangle the effects of memory from those of loser effects (losers behaving subordinately due to prior defeats). Here, we test whether loser effects or individual memory mediate contest behaviour in the African cichlid, Julidochromis transcriptus. We find that on days 3 and 5 after initial contests, losers display subordinate behaviour to contest winners but not to novel contestants. However, this effect disappears after 7 days, at which time losers do not display subordinate behaviour to either rival. These results show that (1) this fish can recall a previously dominant contestant for up to 5 days and (2) as no subordinate displays were shown to the novel contestant, there are no evidences for loser effects in this species. Such short-term memory of past interactions may have broad significance in social species with repeated interactions.
Reddon, Adam R; Gutiérrez-Ibáñez, Cristián; Wylie, Douglas R; Hurd, Peter L
Cerebral lateralization, the partitioning of cognitive tasks to one cerebral hemisphere, is a widespread phenomenon among vertebrates. Despite this diversity, every species studied to date shows substantial individual variation in the strength of lateralization. The neural basis of this trait is unclear, although asymmetries in cerebral structures have been investigated for over a century. The habenular nuclei, for example, have been shown to present striking neuroanatomical and/or neurochemical asymmetries in species ranging from jawless fish to mammals. In teleost fish, these nuclei are relatively symmetrical in most species. Those teleosts that do have asymmetrical habenular nuclei, show varying patterns of asymmetry in different species. Here we investigate the relationship between individual variation of asymmetry in the habenula of a South American cichlid fish, Geophagus brasiliensis, and behaviour in a commonly used test for visual laterality in fish, the detour task. We show that the strength of asymmetry in the habenula is correlated with strength of behavioural lateralization in the detour task. Both the strength and direction of habenular asymmetry are correlated with individual differences in growth rate. We suggest that this relationship results from processes linking growth rate and sexual differentiation to frequency-dependent variation in life-history strategies. To our knowledge, this is the first study to demonstrate a relationship at the individual level between neural asymmetry and lateralized behaviour in a fish.
Wen, C M; Chen, M M; Nan, F H; Wang, C S
In this study, cultures of neural stem-progenitor cells (NSPC) from the brain of green terror cichlid Aequidens rivulatus were established and various NSPCs were demonstrated using immunocytochemistry. All of the NSPCs expressed brain lipid-binding protein, dopamine- and cAMP-regulated neuronal phosphoprotein 32 (DARPP-32), oligodendrocyte transcription factor 2, paired box 6 and sex determining region Y-box 2. The intensity and localisation of these proteins, however, varied among the different NSPCs. Despite being intermediate cells, NSPCs can be divided into radial glial cells, oligodendrocyte progenitor cells (OPC) and neuroblasts by expressing the astrocyte marker glial fibrillary acidic protein (GFAP), OPC marker A2B5 and neuronal markers, including acetyl-tubulin, βIII-tubulin, microtubule-associated protein 2 and neurofilament protein. Nevertheless, astrocytes were polymorphic and were the most dominant cells in the NSPC cultures. By using Matrigel, radial glia exhibiting a long GFAP(+) or DARPP-32(+) fibre and neurons exhibiting a significant acetyl-tubulin(+) process were obtained. The results confirmed that NSPCs obtained from A. rivulatus brains can proliferate and differentiate into neurons in vitro. Clonal culture can be useful for further studying the distinct NSPCs.
Ghosh, Jayati; Kalipeni, Ezekiel
Malawi, a very poor country located in southern Africa, is no exception to the growing trend and severity in HIV prevalence. By the end of 2003 there were 900 000 adults and children in Malawi living with HIV/AIDS. Adult prevalence was estimated to be 15%, which is higher than the 7.1% average rate for sub-Saharan Africa. In order to understand the spread of HIV/AIDS it is imperative to address the economic, social, cultural, and political issues that impact on women's contraction and spread of the virus. We do so in this paper by critically examining the gendered context of HIV/AIDS with reference to Malawi. The theoretical framework for this research focuses on poverty, gender relations, regional migration patterns, and global economic changes which place women in highly vulnerable situations. The study was conducted in a low-income area in Lilongwe, the capital city of Malawi. In 2003 and 2004, 60 randomly selected women who lived in a low socioeconomic residential area completed a structured interview on issues concerning individual economic situations, marriage history, fertility, family planning and social networks, gender, sexual partnerships, and HIV/AIDS. Focus group interviews were also conducted with an additional 20 women. The results of our study indicate that the rising epidemic among women in Malawi is firstly driven by poverty which limits their options. Secondly, gender inequality and asymmetrical sexual relations are basic to spreading HIV/AIDS among women. Thirdly, in spite of their awareness through media and health care professionals, women are unable to protect themselves, which further increases their vulnerability.
Background Shortages of health workers are obstacles to utilising global health initiative (GHI) funds effectively in Africa. This paper reports and analyses two countries' health workforce responses during a period of large increases in GHI funds. Methods Health facility record reviews were conducted in 52 facilities in Malawi and 39 facilities in Zambia in 2006/07 and 2008; quarterly totals from the last quarter of 2005 to the first quarter of 2008 inclusive in Malawi; and annual totals for 2004 to 2007 inclusive in Zambia. Topic-guided interviews were conducted with facility and district managers in both countries, and with health workers in Malawi. Results Facility data confirm significant scale-up in HIV/AIDS service delivery in both countries. In Malawi, this was supported by a large increase in lower trained cadres and only a modest increase in clinical staff numbers. Routine outpatient workload fell in urban facilities, in rural health centres and in facilities not providing antiretroviral treatment (ART), while it increased at district hospitals and in facilities providing ART. In Zambia, total staff and clinical staff numbers stagnated between 2004 and 2007. In rural areas, outpatient workload, which was higher than at urban facilities, increased further. Key informants described the effects of increased workloads in both countries and attributed staff migration from public health facilities to non-government facilities in Zambia to PEPFAR. Conclusions Malawi, which received large levels of GHI funding from only the Global Fund, managed to increase facility staff across all levels of the health system: urban, district and rural health facilities, supported by task-shifting to lower trained staff. The more complex GHI arena in Zambia, where both Global Fund and PEPFAR provided large levels of support, may have undermined a coordinated national workforce response to addressing health worker shortages, leading to a less effective response in rural areas
Berman, Joshua; Mitambo, Collins; Matanje-Mwagomba, Beatrice; Khan, Shiraz; Kachimanga, Chiyembekezo; Wroe, Emily; Mwape, Lonia; van Oosterhout, Joep J; Chindebvu, Getrude; van Schoor, Vanessa; Puchalski Ritchie, Lisa M; Panisset, Ulysses; Kathyola, Damson
With the support of the World Health Organization's Evidence-Informed Policy Network, knowledge translation platforms have been developed throughout Africa, the Americas, Eastern Europe, and Asia to further evidence-informed national health policy. In this commentary, we discuss the approaches, activities and early lessons learned from the development of a Knowledge Translation Platform in Malawi (KTPMalawi). Through ongoing leadership, as well as financial and administrative support, the Malawi Ministry of Health has strongly signalled its intention to utilize a knowledge translation platform methodology to support evidence-informed national health policy. A unique partnership between Dignitas International, a medical and research non-governmental organization, and the Malawi Ministry of Health, has established KTPMalawi to engage national-level policymakers, researchers and implementers in a coordinated approach to the generation and utilization of health-sector research. Utilizing a methodology developed and tested by knowledge translation platforms across Africa, a stakeholder mapping exercise and initial capacity building workshops were undertaken and a multidisciplinary Steering Committee was formed. This Steering Committee prioritized the development of two initial Communities of Practice to (1) improve data utilization in the pharmaceutical supply chain and (2) improve the screening and treatment of hypertension within HIV-infected populations. Each Community of Practice's mandate is to gather and synthesize the best available global and local evidence and produce evidence briefs for policy that have been used as the primary input into structured deliberative dialogues. While a lack of sustained initial funding slowed its early development, KTPMalawi has greatly benefited from extensive technical support and mentorship by an existing network of global knowledge translation platforms. With the continued support of the Malawi Ministry of Health and the
Colbourn, Tim; Lufesi, Norman; Costello, Anthony; Nambiar, Bejoy; Thammasitboon, Satid; Makwenda, Charles; Mwansambo, Charles; McCollum, Eric D.; King, Carina
Background Pneumonia is the leading infectious cause of under-5 mortality in sub-Saharan Africa. Clinical prediction tools may aide case classification, triage, and allocation of hospital resources. We performed an external validation of two published prediction tools and compared this to a locally developed tool to identify children admitted with pneumonia at increased risk for in-hospital mortality in Malawi. Methods We retrospectively analyzed the performance of the Respiratory Index of Severity in Children (RISC) and modified RISC (mRISC) scores in a child pneumonia dataset prospectively collected during routine care at seven hospitals in Malawi between 2011–2014. RISC has both an HIV-infected and HIV-uninfected tool. A local score (RISC-Malawi) was developed using multivariable logistic regression with missing data multiply imputed using chained equations. Score performances were assessed using c-statistics, sensitivity, specificity, positive predictive value, negative predictive value, and likelihood statistics. Results 16,475 in-patient pneumonia episodes were recorded (case-fatality rate (CFR): 3.2%), 9,533 with complete data (CFR: 2.0%). The c-statistic for the RISC (HIV-uninfected) score, used to assess its ability to differentiate between children who survived to discharge and those that died, was 0.72. The RISC-Malawi score, using mid-upper arm circumference as an indicator of malnutrition severity, had a c-statistic of 0.79. We were unable to perform a comprehensive external validation of RISC (HIV-infected) and mRISC as both scores include parameters that were not routinely documented variables in our dataset. Conclusion In our population of Malawian children with WHO-defined pneumonia, the RISC (HIV-uninfected) score identified those at high risk for in-hospital mortality. However the refinement of parameters and resultant creation of RISC-Malawi improved performance. Next steps include prospectively studying both scores to determine if
Barks, Patrick M; Godin, Jean-Guy J
For many animals, the ability to distinguish cues indicative of predation risk from cues unrelated to predation risk is not entirely innate, but rather is learned and improved with experience. Two pathways to such learning are possible. First, an animal could initially express antipredator behaviour toward a wide range of cues and subsequently learn which of those cues are non-threatening. Alternatively, it could initially express no antipredator behaviour toward a wide range of cues and subsequently learn which of them are threatening. While the learned recognition of threatening cues may occur either through personal interaction with a cue (asocial learning) or through observation of the behaviour of social companions toward a cue (social learning), the learned recognition of non-threatening cues seems to occur exclusively through habituation, a form of asocial learning. Here, we tested whether convict cichlid fish (Amatitlaniasiquia) can socially learn to recognize visual cues in their environment as either threatening or non-threatening. We exposed juvenile convict cichlids simultaneously to a novel visual cue and one of three (visual) social cues: a social cue indicative of non-risk (the sight of conspecifics that had previously been habituated to the novel cue), a social cue indicative of predation risk (the sight of conspecifics trained to fear the novel cue), or a control treatment with no social cue. The subsequent response of focal fish, when presented with the novel cue alone, was not influenced by the social cue that they had previously witnessed. We therefore did not find evidence that convict cichlids in our study could use social learning to recognize novel visual cues as either threatening or non-threatening. We consider alternative explanations for our findings.
Hata, Hiroki; Takahashi, Rieko; Ashiwa, Hiroshi; Awata, Satoshi; Takeyama, Tomohiro; Kohda, Masanori; Hori, Michio
The asymmetry of mouth morphology, in which the mouth opens either leftward or rightward, is a unique example of antisymmetry that is widely found in fishes, such as Tanganyikan scale-eating cichlids, herbivorous cichlids, and the Japanese freshwater goby. This dimorphism is thought to be heritable in a Mendelian manner similar to that of the dominance of the lefty allele over righty, with homozygotes of lefty alleles being absent. This study aims to reveal whether this trait is inherited in the same Mendelian manner in fishes other than those examined to date, and whether the absence of the dominant homozygote is due to a lethal effect. We conducted F(1) and F(2) breeding experiments using the Tanganyikan cichlid Julidochromis transcriptus and Japanese medaka Oryzias latipes. For both J. transcriptus and O. latipes, the F(1) generations produced by incrossing lefty parents showed a lefty:righty ratio of 2:1, whereas incrosses of righties produced only righty offspring. Test crosses between lefty and righty produced a 1:1 ratio of lefty and righty offspring. These results are consistent with the inheritance model in which righty is homozygous for a recessive righty allele and lefty is heterozygous for dominant lefty and recessive righty alleles, in agreement with previous observations. The F(2) test cross also confirmed this pattern. Furthermore, no lethal effects on hatchability of O. latipes were detected, refuting the suggestion that the dominant homozygote is embryonic lethal. We conclude that mouth laterality is inherited in the same Mendelian manner in these two species, similar to that other fish species studied previously. However, the reason for absence of the dominant homozygote remains unclear.
Barks, Patrick M.; Godin, Jean-Guy J.
For many animals, the ability to distinguish cues indicative of predation risk from cues unrelated to predation risk is not entirely innate, but rather is learned and improved with experience. Two pathways to such learning are possible. First, an animal could initially express antipredator behaviour toward a wide range of cues and subsequently learn which of those cues are non-threatening. Alternatively, it could initially express no antipredator behaviour toward a wide range of cues and subsequently learn which of them are threatening. While the learned recognition of threatening cues may occur either through personal interaction with a cue (asocial learning) or through observation of the behaviour of social companions toward a cue (social learning), the learned recognition of non-threatening cues seems to occur exclusively through habituation, a form of asocial learning. Here, we tested whether convict cichlid fish (Amatitlaniasiquia) can socially learn to recognize visual cues in their environment as either threatening or non-threatening. We exposed juvenile convict cichlids simultaneously to a novel visual cue and one of three (visual) social cues: a social cue indicative of non-risk (the sight of conspecifics that had previously been habituated to the novel cue), a social cue indicative of predation risk (the sight of conspecifics trained to fear the novel cue), or a control treatment with no social cue. The subsequent response of focal fish, when presented with the novel cue alone, was not influenced by the social cue that they had previously witnessed. We therefore did not find evidence that convict cichlids in our study could use social learning to recognize novel visual cues as either threatening or non-threatening. We consider alternative explanations for our findings. PMID:24086648
2007-01-01The white ring around Lake Powell tells the story. The surface is down 98 feet. This is critical, because Powell, Lake Mead, and other lakes along the Colorado River provide water for millions of people in five states. We are in the eighth year of a drought on the Colorado River. This year was the driest year ever reported in Southern California, and there is a severe drought in Northern California, down to less than 30-percent of snow pack. This ASTER image of part of Lake Powell was acquired in 2001. The gray area depicts the shrunken, reduced 2007 lake extent compared to the extended, larger black area in 2001. The image covers an area of 24 x 30 km, and is centered near 37.1 degrees north latitude, 111.3 degrees west longitude. The U.S. science team is located at NASA's Jet Propulsion Laboratory, Pasadena, Calif. The Terra mission is part of NASA's Science Mission Directorate.
Dobson, J.E.; Peplies, R.W.; Rush, R.M.
This paper examined a National Research Council (NRC) report called Acid Deposition: Long-Term Trends. The report has been the final word on acid deposition as the cause of acidification of lakes. The authors considered it important that the tentative nature of this report be kept in perspective so that the work of the NRC would promote rather than inhibit scientific inquiry on the lake acidification issue. In this spirit, this report proposed that degradation of storm damaged trees could increase the acidity of the forest humus and as a result the ground water which would fed local streams and lakes. They proposed that extensive forest blowdown could be a factor in acidification of surface waters.
Henry, Jaymie Ang; Frenkel, Erica; Borgstein, Eric; Mkandawire, Nyengo; Goddia, Cyril
Background Surgery is increasingly recognized as an important driver for health systems strengthening, especially in developing countries. To facilitate quality improvement initiatives, baseline knowledge of capacity for surgical, anaesthetic, emergency and obstetric care is critical. In partnership with the Malawi Ministry of Health, we quantified government hospitals’ surgical capacity through workforce, infrastructure and health service delivery components. Methods From November 2012 to January 2013, we surveyed district and mission hospital administrators and clinical staff onsite using a modified version of the Personnel, Infrastructure, Procedures, Equipment and Supplies (PIPES) tool from Surgeons OverSeas. We calculated percentage of facilities demonstrating adequacy of the assessed components, surgical case rates, operating theatre density and surgical workforce density. Results Twenty-seven government hospitals were surveyed (90% of the district hospitals, all central hospitals). Of the surgical workforce surveyed (n = 370), 92.7% were non-surgeons and 77% were clinical officers (COs). Of the 109 anaesthesia providers, 95.4% were non-physician anaesthetists (anaesthesia COs or ACOs). Non-surgeons and ACOs were the only providers of surgical services and anaesthetic services in 85% and 88.9% of hospitals, respectively. No specialists served the district hospitals. All of the hospitals experienced periods without external electricity. Most did not always have a functioning generator (78.3% district, 25% central) or running water (82.6%, 50%). None of the district hospitals had an Intensive Care Unit (ICU). Cricothyroidotomy, bowel resection and cholecystectomy were not done in over two-thirds of hospitals. Every hospital provided general anaesthesia but some did not always have a functioning anaesthesia machine (52.2%, 50%). Surgical rate, operating theatre density and surgical workforce density per 100 000 population was 289.48–747.38 procedures, 0
Glynn, Judith R.; Kayuni, Ndoliwe; Floyd, Sian; Banda, Emmanuel; Francis-Chizororo, Monica; Tanton, Clare; Molesworth, Anna; Hemmings, Joanne; Crampin, Amelia C.; French, Neil
Background Age at sexual debut is a key behavioural indicator used in HIV behavioural surveillance. Early age at menarche may precipitate early sex through perceived readiness for sex, or through school drop-out, but this is rarely studied. We investigated trends and circumstances of sexual debut in relation to schooling and age at menarche. Methods and Findings A cross-sectional sexual behaviour survey was conducted on all individuals age 15–59 within a demographic surveillance site in Karonga District, Malawi. Time trends were assessed using birth cohorts. Survival analysis was used to estimate the median age at menarche, sexual debut and first marriage. The 25th centile was used to define “early” sex, and analyses of risk factors for early sex were restricted to those who had reached that age, and were done using logistic regression. Of the 8232 women and 7338 men resident in the area, 88% and 78%, respectively, were seen, and, 94% and 92% of these were interviewed. The median reported age at first sex was 17.5 for women and 18.8 for men. For women, ages at menarche, sexual debut and first marriage did not differ by birth cohort. For men, age at sexual debut and first marriage decreased slightly in later birth cohorts. For both men and women increased schooling was associated with later sexual debut and a longer delay between sexual debut and first marriage, but the associations were stronger for women. Earlier age at menarche was strongly associated with earlier sexual debut and marriage and lower schooling levels. In women early sexual debut (<16 years) was less likely in those with menarche at age 14–15 (odds ratio (OR) 0.31, 95%CI 0.26–0.36), and ≥16 (OR 0.04, 95%CI 0.02–0.05) compared to those with menarche at <14. The proportion of women who completed primary school was 46% in those with menarche at <14, 60% in those with menarche at 14–15 and 70% in those with menarche at ≥16. The association between age at menarche and schooling was
Hrapcak, Susan; Kuper, Hannah; Bartlett, Peter; Devendra, Akash; Makawa, Atupele; Kim, Maria; Kazembe, Peter; Ahmed, Saeed
Introduction With improved access to antiretroviral therapy (ART), HIV infection is becoming a chronic illness. Preliminary data suggest that HIV-infected children have a higher risk of disabilities, including hearing impairment, although data are sparse. This study aimed to estimate the prevalence and types of hearing loss in HIV-infected children in Lilongwe, Malawi. Methods This was a cross-sectional survey of 380 HIV-infected children aged 4–14 years attending ART clinic in Lilongwe between December 2013-March 2014. Data was collected through pediatric quality of life and sociodemographic questionnaires, electronic medical record review, and detailed audiologic testing. Hearing loss was defined as >20 decibels hearing level (dBHL) in either ear. Predictors of hearing loss were explored by regression analysis generating age- and sex-adjusted odds ratios. Children with significant hearing loss were fitted with hearing aids. Results Of 380 patients, 24% had hearing loss: 82% conductive, 14% sensorineural, and 4% mixed. Twenty-one patients (23% of those with hearing loss) were referred for hearing aid fitting. There was a higher prevalence of hearing loss in children with history of frequent ear infections (OR 7.4, 4.2–13.0) and ear drainage (OR 6.4, 3.6–11.6). Hearing loss was linked to history of WHO Stage 3 (OR 2.4, 1.2–4.5) or Stage 4 (OR 6.4, 2.7–15.2) and history of malnutrition (OR 2.1, 1.3–3.5), but not to duration of ART or CD4. Only 40% of caregivers accurately perceived their child’s hearing loss. Children with hearing impairment were less likely to attend school and had poorer emotional (p = 0.02) and school functioning (p = 0.04). Conclusions There is an urgent need for improved screening tools, identification and treatment of hearing problems in HIV-infected children, as hearing loss was common in this group and affected school functioning and quality of life. Clear strategies were identified for prevention and treatment, since most
Gilbert, Grove Karl
This volume is a contribution to the later physical history of the Great Basin. As a geographic province the Great Basin is characterized by a dry climate, changes of drainage, volcanic eruption, and crustal displacement. Lake Bonneville, the special theme of the volume, was a phenomenon of climate and drainage, but its complete history includes an account of contemporaneous eruption and displacement.
van Dongen, Wouter F D; Wagner, Richard H; Moodley, Yoshan; Schaedelin, Franziska C
Animal dispersal is associated with diverse costs and benefits that vary among individuals based on phenotype and ecological conditions. For example, females may disperse when males benefit more from defending territories in familiar environments. Similarly, size differences in dispersal propensity may occur when dispersal costs are size-dependent. When individuals do disperse, they may adopt behavioral strategies that minimize dispersal costs. Dispersing fish, for example, may travel within shoals to reduce predation risks. Further, kin shoaling may augment inclusive fitness by reducing predation of relatives. However, studies are lacking on the role of kin shoaling in dispersal. We explored how sex and size influence dispersal and kin shoaling in the cichlid Neolamprologus caudopunctatus. We microsatellite genotyped over 900 individuals from two populations separated by a potential dispersal barrier, and documented patterns of population structure, migration and within-shoal relatedness. Genetic differentiation across the barrier was greater for smaller than larger fish, suggesting larger fish had dispersed longer distances. Females exhibited weaker genetic differentiation and 11 times higher migration rates than males, indicating longer-distance female-biased dispersal. Small females frequently shoaled with siblings, possibly offsetting dispersal costs associated with higher predation risks. In contrast, small males appeared to avoid kin shoaling, possibly to avoid local resource competition. In summary, long-distance dispersal in N. caudopunctatus appears to be female-biased, and kin-based shoaling by small females may represent a behavioral adaptation that reduces dispersal costs. Our study appears to be the first to provide evidence that sex differences in dispersal influence sex differences in kin shoaling.
Becker, Lisa; Hilliard, Austin T.; Kent, Kai R.; Fernald, Russell D.
ABSTRACT Male African cichlid fish, Astatotilapia burtoni, have been classified as dominant or subordinate, each with unique behavioral and endocrine profiles. Here we characterize two distinct subclasses of dominant males based on types of aggressive behavior: (1) males that display escalating levels of aggression and court females while they establish a territory, and (2) males that display a stable level of aggression and delay courting females until they have established a territory. To profile differences in their approach to a challenge, we used an intruder assay. In every case, there was a male-male confrontation between the resident dominant male and the intruder, with the intruder quickly taking a subordinate role. However, we found that dominant males with escalating aggression spent measurably more time attacking subordinates than did dominant males with stable aggression that instead increased their attention toward the females in their tank. There was no difference in the behavior of intruders exposed to either type of dominant male, suggesting that escalating aggression is an intrinsic characteristic of some dominant males and is not elicited by the behavior of their challengers. Male behavior during the first 15 min of establishing a territory predicts their aggressive class. These two types of dominant males also showed distinctive physiological characteristics. After the intruder assay, males with escalating aggression had elevated levels of 11-ketotestosterone (11-KT), testosterone, estradiol, and cortisol, while those with stable aggression did not. These observations show that the same stimulus can elicit different behavioral and endocrine responses among A. burtoni dominant males that characterize them as either escalating or stable aggressive types. Our ability to identify which individuals within a population have escalating levels of aggressive responses versus those which have stable levels of aggressive responses when exposed to the same
Heg, Dik; Taborsky, Michael
Background We manipulated predation risk in a field experiment with the cooperatively breeding cichlid Neolamprologus pulcher by releasing no predator, a medium- or a large-sized fish predator inside underwater cages enclosing two to three natural groups. We assessed whether helpers changed their helping behaviour, and whether within-group conflict changed, depending on these treatments, testing three hypotheses: ‘pay-to-stay’ PS, ‘risk avoidance’ RA, or (future) reproductive benefits RB. We also assessed whether helper food intake was reduced under risk, because this might reduce investments in other behaviours to save energy. Methodology/Principal Findings Medium and large helpers fed less under predation risk. Despite this effect helpers invested more in territory defence, but not territory maintenance, under the risk of predation (supporting PS). Experimentally covering only the breeding shelter with sand induced more helper digging under predation risk compared to the control treatment (supporting PS). Aggression towards the introduced predator did not differ between the two predator treatments and increased with group member size and group size (supporting PS and RA). Large helpers increased their help ratio (helping effort/breeder aggression received, ‘punishment’ by the dominant pair in the group) in the predation treatments compared to the control treatment, suggesting they were more willing to PS. Medium helpers did not show such effects. Large helpers also showed a higher submission ratio (submission/ breeder aggression received) in all treatments, compared to the medium helpers (supporting PS). Conclusions/Significance We conclude that predation risk reduces helper food intake, but despite this effect, helpers were more willing to support the breeders, supporting PS. Effects of breeder punishment suggests that PS might be more important for large compared to the medium helpers. Evidence for RA was also detected. Finally, the results were
Hesse, Saskia; Thünken, Timo
Living in groups is a widespread phenomenon in many animal taxa. The reduction of predation risk is thought to be an important cause for the formation of groups. Consequently, grouping behavior is particularly pronounced during vulnerable life stages, i.e., as juveniles. However, group living does not only provide benefits but also imposes costs on group members, e.g., increased competition for food. Thus, benefits of grouping behavior might not be evident when predation risk is absent. The adaptive significance of living and also developing in a group independent from predation risk has received relatively little attention although this might have important implications on the evolution and maintenance of group living. The first aim of the present study was to examine whether the social environment affects juvenile performance in the cichlid fish Pelvicachromis taeniatus and, secondly, whether kinship affects social behavior. Kin selection theory predicts benefits from grouping with kin. Here, we demonstrate that juveniles reared in a group grow on average faster compared to juveniles reared in isolation under standardized laboratory conditions without predation risk. Furthermore, we found significant differences in social behavior between juveniles reared in a group and reared in isolation. Fish reared in isolation were significantly more aggressive and less willing to shoal than group-reared fish. As expected, genetic relatedness influenced social behavior in group-reared fish as well: dyads of juveniles consisting of kin showed increased group cohesiveness compared to non-kin dyads. We discuss the potential benefits of group living in general and living with kin in particular.
Almeida, Olinda; Gonçalves-de-Freitas, Eliane; Lopes, João S; Oliveira, Rui F
Androgens are known to respond to social challenges and to control the expression of social behavior and reproductive traits, such as gonadal maturation and sperm production, expression of secondary sex characters and reproductive behaviors. According to the challenge hypothesis variation in androgen levels above a breeding baseline should be explained by the regime of social challenges faced by the individual considering the trade-offs of androgens with other traits (e.g. parental care). One prediction that can be derived from the challenge hypothesis is that androgen levels should increase in response to social instability. Moreover, considering that a tighter association of relevant traits is expected in periods of environmental instability, we also predict that in unstable environments the degree of correlations among different behaviors should increase and hormones and behavior should be associated. These predictions were tested in a polygamous cichlid fish (Mozambique tilapia, Oreochromis mossambicus) with exclusive maternal care. Social instability was produced by swapping dominant males among groups. Stable treatment consisted in removing and placing back dominant males in the same group, in order to control for handling stress. Cortisol levels were also measured to monitor stress levels involved in the procedure and their relation to the androgen patterns and behavior. As predicted androgen levels increased in males in response to the establishment of a social hierarchy and presence of receptive females. However, there were no further differential increases in androgen levels over the social manipulation phase between social stable and social unstable groups. As predicted behaviors were significantly more correlated among themselves in the unstable than in the stable treatment and an associated hormone-behavior pattern was only observed in the unstable treatment.
Hesse, Saskia; Thünken, Timo
Living in groups is a widespread phenomenon in many animal taxa. The reduction of predation risk is thought to be an important cause for the formation of groups. Consequently, grouping behavior is particularly pronounced during vulnerable life stages, i.e., as juveniles. However, group living does not only provide benefits but also imposes costs on group members, e.g., increased competition for food. Thus, benefits of grouping behavior might not be evident when predation risk is absent. The adaptive significance of living and also developing in a group independent from predation risk has received relatively little attention although this might have important implications on the evolution and maintenance of group living. The first aim of the present study was to examine whether the social environment affects juvenile performance in the cichlid fish Pelvicachromis taeniatus and, secondly, whether kinship affects social behavior. Kin selection theory predicts benefits from grouping with kin. Here, we demonstrate that juveniles reared in a group grow on average faster compared to juveniles reared in isolation under standardized laboratory conditions without predation risk. Furthermore, we found significant differences in social behavior between juveniles reared in a group and reared in isolation. Fish reared in isolation were significantly more aggressive and less willing to shoal than group-reared fish. As expected, genetic relatedness influenced social behavior in group-reared fish as well: dyads of juveniles consisting of kin showed increased group cohesiveness compared to non-kin dyads. We discuss the potential benefits of group living in general and living with kin in particular.
Nolan, Brian C
Laboratory activities serve several important functions in undergraduate science education. For neuroscience majors, an important and sometimes underemphasized tool is the use of behavioral observations to help inform us about the consequences of changes that are occurring on a neuronal level. To help address this concern, the following laboratory exercise is presented. The current project tested the prediction that the most dominant fish in a tank of cichlids will have gained the most benefits of its position resulting in the greatest growth and hence, become the largest fish. More specifically: (1) is there evidence that a social hierarchy exists among the fish in our tank based on the number of aggressive acts among the four largest fish; (2) if so, does the apparent rank correspond to the size of the fish as predicted by previous studies? Focal sampling and behavior sampling of aggressive acts between fish were utilized in the data collection. Collectively, the data suggest a social dominance hierarchy may be in place with the following rank order from highest to lowest: Fish A > Fish B > Fish D > Fish C. While the largest (Fish A) seems to be at the top, Fish C ended up being ranked lower than Fish D despite the fact that Fish C is larger. Overall, the project was considered a success by the instructor and students. The students offered several suggestions that could improve future versions of this type of project, in particular concerning the process of constructing a poster about the project. The implications of the data and student learning outcomes are discussed.
Maruska, Karen P; Fernald, Russell D
Social interactions with the outcome of a position in a dominance hierarchy can have profound effects on reproductive behavior and physiology, requiring animals to integrate environmental information with their internal physiological state; but how is salient information from the animal's dynamic social environment transformed into adaptive behavioral, physiological, and molecular-level changes? The African cichlid fish, Astatotilapia burtoni, is ideally suited to understand socially controlled reproductive plasticity because activity of the male reproductive (brain-pituitary-gonad) axis is tightly linked to social status. Males form hierarchies in which a small percentage of brightly colored dominant individuals have an active reproductive axis, defend territories, and spawn with females, while the remaining males are subordinate, drably colored, do not hold a territory, and have a suppressed reproductive system with minimal opportunities for spawning. These social phenotypes are plastic and quickly reversible, meaning that individual males may switch between dominant and subordinate status multiple times within a lifetime. Here, we review the rapid and remarkable plasticity that occurs along the entire reproductive axis when males rise in social rank, a transition that has important implications for the operational sex ratio of the population. When males rise in rank, transformations occur in the brain, pituitary, circulation, and testes over short time-scales (minutes to days). Changes are evident in overt behavior, as well as modifications at the physiological, cellular, and molecular levels that regulate reproductive capacity. Widespread changes triggered by a switch in rank highlight the significance of external social information in shaping internal physiology and reproductive competence.
Raffini, Francesca; Fruciano, Carmelo; Franchini, Paolo; Meyer, Axel
How polymorphisms consisting in left-right asymmetries are produced and maintained in natural populations is a tantalizing question, which remains largely unanswered. The scale-eating cichlid fish Perissodus microlepis is a remarkable example of extreme ecological specialization achieved by morphological and behavioural laterality. Its asymmetric mouth is accompanied by a pronounced lateralized foraging behaviour, where a left-bending morph preferentially feeds on the scales of the right side of its prey, while the opposite is true for the right morph. This striking asymmetry made this fish a textbook example of the astounding degree of ecological specialization and negative frequency-dependent selection. Yet, the genetic basis underlying this spectacular laterality remains unknown. We addressed this question through analyses of wild-caught fish using high-throughput DNA sequencing data. A novel array of SNP markers was developed by ddRAD sequencing (ddRADseq) and the use of pooled DNA samples (PoolSeq). We obtained more than 155 000 SNPs using ddRADseq and 3 900 000 SNPs with PoolSeq. Among these, we identified one (ddRAD) SNP, and 38 or 378 (PoolSeq) windows that are differentiated between the left and right morphs accounting for spurious associations due to geographic structuring. This allowed us to uncover candidate genomic regions that potentially contain genes for this trait. Then, this interesting trait has a genetic basis that is likely to be influenced by multiple loci. This result contributes to a greater understanding of the genetic bases of left-right asymmetry and, ultimately, the evolutionary processes governing the maintenance of this striking case of laterality.
HARBOTT, LENE K.; BURMEISTER, SABRINA S.; WHITE, RICHARD B.; VAGELL, MIKE; FERNALD, RUSSELL D.
Androgens are an important output of the hypothalamic-pituitary-gonadal (HPG) axis that controls reproduction in all vertebrates. In male teleosts two androgens, testosterone and 11-ketotestosterone, control sexual differentiation and development in juveniles and reproductive behavior in adults. Androgenic signals provide feedback at many levels of the HPG axis, including the hypothalamic neurons that synthesize and release gonadotropin-releasing hormone 1 (GnRH1), but the precise cellular site of androgen action in the brain is not known. Here we describe two androgen receptor subtypes, ARα and ARβ, in the cichlid Astatotilapia burtoni and show that these subtypes are differentially located throughout the adult brain in nuclei known to function in the control of reproduction. ARα was expressed in the ventral part of the ventral telencephalon, the preoptic area (POA) of the hypothalamus and the ventral hypothalamus, whereas ARβ was more widely expressed in the dorsal and ventral telencephalon, the POA, and the ventral and dorsal hypothalamus. We provide the first evidence in any vertebrate that the GnRH1-releasing neurons, which serve as the central control point of the HPG axis, express both subtypes of AR. Using quantitative real-time PCR, we show that A. burtoni AR subtypes have different expression levels in adult tissue, with ARα showing significantly higher expression than ARβ in the pituitary, and ARβ expressed at a higher level than ARα in the anterior and middle brain. These data provide important insight into the role of androgens in regulating the vertebrate reproductive axis. PMID:17614300
Schoenleber, J.; Anken, R.
Previous investigations revealed that the growth of fish inner ear otoliths depends on the amplitude and direction of gravity, thus suggesting the existence of a (negative) feedback mechanism. In the course of these experiments, it was shown that altered gravity both affected otolith size (and thus the provision of the proteinacious matrix) as well as the incorporation of calcium. It is hitherto unknown, as of whether sensory hair cells are involved either in the regulation of otolith growth or in the provision of otolithic material (such as protein or inorganic components) or even both. Developing cichlid fish Oreochromis mossambicus were therefore immersed in varying concentrations of the ototoxic aminoglycoside gentamicin (20, 40, 60, 100, 120mg/l; at even higher concentrations, the animals did not survive) for 10 or 20 days. At the beginning and at the end of the experimental periods, the fish were incubated in the calcium-tracer alizarin complexone (AC). After the experiment, otoliths were dissected and the area grown during gentamicin exposure (i.e., the area enclosed by the two AC labellings) was determined planimetrically. The results clearly showed that incubating the animals in a gentamicin solution had no effect on otolith growth. Further examinations, focused on the ultrastructure and on physiological parameters of the sensory hair cells, revealed, that these cells had not been affected by gentamicin treatment. These findings suggest that gentamicin does either not act ototoxically on hair cells in developing fish (in contrast to adult fish and developing mammals), or that immersion in gentamicin does not allow the aminoglycoside to act on inner ear hair cells (although it damages lateral line neuromasts and high concentrations affect the rate of survival). The role of sensory cells on the formation of otoliths thus remains obscure. Acknowledgement: This work was financially supported by the German Aerospace Center (DLR) (FKZ: 50 WB 9997).
Kültz, Dietmar; Li, Johnathon; Gardell, Alison; Sacchi, Romina
A two-tiered label-free quantitative (LFQ) proteomics workflow was used to elucidate how salinity affects the molecular phenotype, i.e. proteome, of gills from a cichlid fish, the euryhaline tilapia (Oreochromis mossambicus). The workflow consists of initial global profiling of relative tryptic peptide abundances in treated versus control samples followed by targeted identification (by MS/MS) and quantitation (by chromatographic peak area integration) of validated peptides for each protein of interest. Fresh water acclimated tilapia were independently exposed in separate experiments to acute short-term (34 ppt) and gradual long-term (70 ppt, 90 ppt) salinity stress followed by molecular phenotyping of the gill proteome. The severity of salinity stress can be deduced with high technical reproducibility from the initial global label-free quantitative profiling step alone at both peptide and protein levels. However, an accurate regulation ratio can only be determined by targeted label-free quantitative profiling because not all peptides used for protein identification are also valid for quantitation. Of the three salinity challenges, gradual acclimation to 90 ppt has the most pronounced effect on gill molecular phenotype. Known salinity effects on tilapia gills, including an increase in the size and number of mitochondria-rich ionocytes, activities of specific ion transporters, and induction of specific molecular chaperones are reflected in the regulation of abundances of the corresponding proteins. Moreover, specific protein isoforms that are responsive to environmental salinity change are resolved and it is revealed that salinity effects on the mitochondrial proteome are nonuniform. Furthermore, protein NDRG1 has been identified as a novel key component of molecular phenotype restructuring during salinity-induced gill remodeling. In conclusion, besides confirming known effects of salinity on gills of euryhaline fish, molecular phenotyping reveals novel insight into
Beier, M.; Anken, R.; Rahmann, H.
It has been shown earlier that hypergravity slows down inner ear otolith growth in developing fish. Otolith growth in terms of mineralisation mainly depends on the enzyme carboanhydrase (CAH), which is responsible for the provision of the pH- value necessary for calcium carbonate deposition and thus also is presumed to play a prominent role in Ménière's disease (a sensory - motor disorder inducing vertigo and kinetosis). Larval siblings of cichlid fish (Oreochromis mossambicus) were subjected to hypergravity (3g; 6 hours) during development and separated into normally and kinetotically swimming individuals following the transfer to 1g (i.e., stopping the centrifuge; kinetotically behaving fish performed spinning movements). Subsequently, CAH was histochemically demonstrated in inner ear ionocytes (cells involved in the endolymphatic ion exchange) and enzyme reactivity was determined densitometrically. The results showed that CAH-reactivity was significantly increased in normally behaving hyper-g specimens as compared to controls kept at 1g, whereas no difference in enzyme reactivity was evident between the controls and kinetotically behaving fish. On the background of earlier studies, according to which (1) hypergravity induces a decrease of otolith growth and (2) the otolithic calcium incorporation (visualized using the calcium -tracer alizarin complexone) of kinetotically swimming hyper - g fish was lower as compared to normally behaving hyper - g animals, the present study strongly supports the concept that an increase in CAH-reactivity may result in a decrease of otolithic calcium deposition. The mechanism regulating CAH-activity hitherto remains to be determined. Acknowledgement: This work was financially supported by the German Aerospace Center (DLR) (FKZ: 50 WB 9997).
Schönleber, J.; Anken, R. H.
Previous investigations revealed that the growth of fish inner ear otoliths depends on the amplitude and the direction of gravity, thus suggesting the existence of a (negative) feedback mechanism. In the course of these experiments, it was shown that altered gravity both affected otolith size (and thus the provision of the proteinacious matrix) as well as the incorporation of calcium. It is hitherto unknown, as of whether sensory hair cells are involved either in the regulation of otolith growth or in the provision of otolithic material (such as protein or inorganic components) or even both. The ototoxic aminoglycoside gentamicin (GM) damages hair cells in many vertebrates (and is therefore used for the treatment of Meniere's disease in humans). The present study was thus designed to determine as of whether vestibular sensory cells are needed for otolith growth by applying GM in order to induce a (functionally relevant) loss of these cells. Developing cichlid fish Oreochromis mossambicus were therefore immersed in 120 mg/l GM for 10 or 21 days. At the beginning and at the end of the experimental periods, the fish were incubated in the calcium-tracer alizarin complexone (AC). After the experiment, otoliths were dissected and the area grown during GM-exposure (i.e., the area enclosed by the two AC labellings) was determined planimetrically. The results showed that incubating the animals in a GM-solution had no effect on otolith growth, but the development of otolith asymmetry was affected. Ultrastructural examinations of the sensory hair cells revealed that they had obviously not been affected by GM-treatment (no degenerative morphological features observed). Overall, the present results suggest that hair cells are not affected by GM concerning their possible role in (general) otolith growth, but that these cells indeed might have transitionally been impaired by GM resulting in a decreased capacity of regulating otolith symmetry.
Chen, Chun-Chun; Fernald, Russell D.
Social behavior can influence physiological systems dramatically yet the sensory cues responsible are not well understood. Behavior of male African cichlid fish, Astatotilapia burtoni, in their natural habitat suggests that visual cues from conspecifics contribute significantly to regulation of social behavior. Using a novel paradigm, we asked whether visual cues alone from a larger conspecific male could influence behavior, reproductive physiology and the physiological stress response of a smaller male. Here we show that just seeing a larger, threatening male through a clear barrier can suppress dominant behavior of a smaller male for up to 7 days. Smaller dominant males being “attacked” visually by larger dominant males through a clear barrier also showed physiological changes for up to 3 days, including up-regulation of reproductive- and stress-related gene expression levels and lowered plasma 11-ketotestesterone concentrations as compared to control animals. The smaller males modified their appearance to match that of non-dominant males when exposed to a larger male but they maintained a physiological phenotype similar to that of a dominant male. After 7 days, reproductive- and stress- related gene expression, circulating hormone levels, and gonad size in the smaller males showed no difference from the control group suggesting that the smaller male habituated to the visual intruder. However, the smaller male continued to display subordinate behaviors and assumed the appearance of a subordinate male for a full week despite his dominant male physiology. These data suggest that seeing a larger male alone can regulate the behavior of a smaller male but that ongoing reproductive inhibition depends on additional sensory cues. Perhaps, while experiencing visual social stressors, the smaller male uses an opportunistic strategy, acting like a subordinate male while maintaining the physiology of a dominant male. PMID:21633515
Fernald, Russell D.
In many vertebrates, reproduction is regulated by social interactions in which dominant males control access to females and food. Subordinate males that displace dominant individuals must rapidly adopt behavioral and physiological traits of the higher rank to gain reproductive success. To understand the process of phenotypic plasticity during social ascent, we analyzed the temporal expression pattern of dominance behaviors and circulating androgen levels when socially-suppressed males of an African cichlid fish Astatotilapia burtoni ascended in status. These experiments tested a prediction of the ‘challenge hypothesis’ that, during periods of social instability, male androgen levels are higher than during socially stable times. We found that socially and reproductively suppressed males perform territorial and reproductive behaviors within minutes of an opportunity to ascend in status, and that animals switch from initial expression of territorial behaviors to more reproductive behaviors during territory establishment. Following this rapid response, social stability may be achieved within 1–3 days of social ascent. Consistent with predictions of the ‘challenge hypothesis’, circulating 11-ketotestosterone (11-KT) levels were elevated within 30 min following social opportunity, coincident with increased aggressive behavior. However, territorial behaviors and serum 11-KT levels were then dissociated by 72 hrs after social ascent, suggesting either rapid social stability and/or increase