A genetic demographic analysis of Lake Malawi rock‐dwelling cichlids using spatio‐temporal sampling

We estimated the effective population sizes (N_e) and tested for short‐term temporal demographic stability of populations of two Lake Malawi cichlids: Maylandia benetos, a micro‐endemic, and Maylandia zebra, a widespread species found across the lake. We sampled a total of 351 individuals, genotyped them at 13 microsatellite loci and sequenced their mitochondrial D‐loop to estimate genetic diversity, population structure, demographic history and effective population sizes. At the microsatellite loci, genetic diversity was high in all populations. Yet, genetic diversity was relatively low for the sequence data. Microsatellites yielded mean N_e estimates of 481 individuals (±99 SD) for M. benetos and between 597 (±106.3 SD) and 1524 (±483.9 SD) individuals for local populations of M. zebra. The microsatellite data indicated no deviations from mutation–drift equilibrium. Maylandia zebra was further found to be in migration–drift equilibrium. Temporal fluctuations in allele frequencies were limited across the sampling period for both species. Bayesian Skyline analyses suggested a recent expansion of M. zebra populations in line with lake‐level fluctuations, whereas the demographic history of M. benetos could only be estimated for the very recent past. Divergence time estimates placed the origin of M. benetos within the last 100 ka after the refilling of the lake and suggested that it split off the sympatric M. zebra population. Overall, our data indicate that micro‐endemics and populations in less favourable habitats have smaller N_e, indicating that drift may play an important role driving their divergence. Yet, despite small population sizes, high genetic variation can be maintained.     

Aggression in closely related Malawi cichlids varies inversely with habitat complexity

Within the past 2 million years, the cichlids of Lake Malawi have diversified into well over 500 species resulting in one of the worlds largest lacustrine fish radiations. As a result, many of the habitats within the lake support a high diversity of species. In these highly species rich communities, male cichlids must acquire and defend a territory to successfully reproduce. Within the rock-dwelling cichlids of Lake Malawi (mbuna), this has resulted in the formation of polyspecific leks on the heterogeneous rocky benthos. Aggression is fairly common in these leks and has been tied not only to individual reproduction but to the larger phenomenon of community assembly and the maintenance of biological diversity. In this study, I examined the patterns of aggressive acts of four species within the mbuna genus Maylandia at two locations in the southern Lake Malawi. The number of aggressive acts of two sympatric species was examined at each location. At each site, one species defends territories over bedrock and the other over cobble. The number of aggressive acts across the four species was compared. The influence of habitat type on male aggression was examined and the targets of male aggression were identified to evaluate several hypotheses concerning the evolution of male aggression. The results show that aggression quantitatively varied among species, was largely directed towards heterospecifics, and was strongly influenced by habitat type. The aggressive behavior of one sympatric species pair, Maylandia benetos and Maylandia zebra, was observed under controlled laboratory conditions. Laboratory results support field observations: the bedrock associated species performed more aggressive acts and aggressive acts were directed equally at con- and heterospecifics. The results of this study suggest that habitat complexity plays a larger role in shaping aggressive behavior than other suggested factors such as competition for resources.     

Evolution of body shape in differently coloured sympatric congeners and allopatric populations of Lake Malawi's rock‐dwelling cichlids

The cichlid fishes of Lake Malawi represent one of the most diverse adaptive radiations of vertebrates known. Among the rock‐dwelling cichlids (mbuna), closely related sympatric congeners possess similar trophic morphologies (i.e. cranial and jaw structures), defend overlapping or adjacent territories, but can be easily distinguished based on male nuptial coloration. The apparent morphological similarity of congeners, however, leads to an ecological conundrum: theory predicts that ecological competition should lead to competitive exclusion. Hence, we hypothesized that slight, yet significant, ecological differences accompanied the divergence in sexual signals and that the divergence of ecological and sexual traits is correlated. To evaluate this hypothesis, we quantified body shape, a trait of known ecological importance, in populations of Maylandia zebra, a barred, widespread mbuna, and several sympatric nonbarred congeners. We found that the barred populations differ in body shape from their nonbarred sympatric congeners and that the direction of shape differences was consistent across all barred vs. nonbarred comparisons. Barred populations are generally deeper bodied which may be an adaptation to the structurally complex habitat they prefer, whereas the nonbarred species have a more fusiform body shape, which may be adaptive in their more open microhabitat. Furthermore, M. zebra populations sympatric with nonbarred congeners differ from populations where the nonbarred phenotype is absent and occupy less morphospace, indicating potential ecological character displacement. Mitochondrial DNA as well as published AFLP data indicated that the nonbarred populations are not monophyletic and therefore may have evolved multiple times independently. Overall our data suggest that the evolution of coloration and body shape may be coupled as a result of correlational selection. We hypothesize that correlated evolution of sexually selected and ecological traits may have contributed to rapid speciation as well as the maintenance of diversity in one of the most diverse adaptive radiations known.     

Signal and preference divergence among populations of the non‐endemic basal Lake Malawi cichlid fish Astatotilapia calliptera (Perciformes: Cichlidae)

In Lake Malawi and Lake Victoria, cichlid fishes have diversified into hundreds of species, many reproductively isolated by mate choice. Territorial males tend to be more aggressive to similar‐coloured males, facilitating coexistence of divergent colour morphs or species. Behavioural mate choice and aggression biases of species and allopatric populations of specialized rocky shore cichlids are influenced by divergent signals such as male colour. Believed to be basal to the Lake Malawi haplochromine radiation, and inhabiting shallow weedy areas of the lake and neighbouring water bodies, Astatotilapia calliptera also shows allopatric variation in colour. Here, it is demonstrated that such signal divergence is associated with tendencies of females to mate with males of their own population and also for males to preferentially attack males of their own population, indicating that preference divergence related to signal divergence in allopatry may have operated throughout the adaptive radiation of the Malawian cichlids. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 180–188.     

Evolutionary divergence in life history traits among populations of the Lake Malawi cichlid fish Astatotilapia calliptera

During the early stages of adaptive radiation, populations diverge in life history traits such as egg size and growth rates, in addition to eco‐morphological and behavioral characteristics. However, there are few studies of life history divergence within ongoing adaptive radiations. Here, we studied Astatotilapia calliptera, a maternal mouthbrooding cichlid fish within the Lake Malawi haplochromine radiation. This species occupies a rich diversity of habitats, including the main body of Lake Malawi, as well as peripheral rivers and shallow lakes. We used common garden experiments to test for life history divergence among populations, focussing on clutch size, duration of incubation, egg mass, offspring size, and growth rates. In a first experiment, we found significant differences among populations in average clutch size and egg mass, and larger clutches were associated with smaller eggs. In a second experiment, we found significant differences among populations in brood size, duration of incubation, juvenile length when released, and growth rates. Larger broods were associated with smaller juveniles when released and shorter incubation times. Although juvenile growth rates differed between populations, these were not strongly related to initial size on release. Overall, differences in life history characters among populations were not predicted by major habitat classifications (Lake Malawi or peripheral habitats) or population genetic divergence (microsatellite‐based F_ST). We suggest that the observed patterns are consistent with local selective forces driving the observed patterns of trait divergence. The results provide strong evidence of evolutionary divergence and covariance of life history traits among populations within a radiating cichlid species, highlighting opportunities for further work to identify the processes driving the observed divergence.     

Geography disentangles introgression from ancestral polymorphism in Lake Malawi cichlids

Phenotypically diverse Lake Malawi cichlids exhibit similar genomes. The extensive sharing of genetic polymorphism among forms has both intrigued and frustrated biologists trying to understand the nature of diversity in this and other rapidly evolving systems. Shared polymorphism might result from hybridization and/or the retention of ancestrally polymorphic alleles. To examine these alternatives, we used new genomic tools to characterize genetic differentiation in widespread, geographically structured populations of Labeotropheus fuelleborni and Metriaclima zebra. These phenotypically distinct species share mitochondrial DNA (mtDNA) haplotypes and show greater mtDNA differentiation among localities than between species. However, Bayesian analysis of nuclear single nucleotide polymorphism (SNP) data revealed two distinct genetic clusters corresponding perfectly to morphologically diagnosed L. fuelleborni and M. zebra. This result is a function of the resolving power of the multi‐locus dataset, not a conflict between nuclear and mitochondrial partitions. Locus‐by‐locus analysis showed that mtDNA differentiation between species (F_CT) was nearly identical to the median single‐locus SNP F_CT. Finally, we asked whether there is evidence for gene flow at sites of co‐occurrence. We used simulations to generate a null distribution for the level of differentiation between co‐occurring populations of L. fuelleborni and M. zebra expected if there was no hybridization. The null hypothesis was rejected for the SNP data; populations that co‐occur at rock reef sites were slightly more similar than expected by chance, suggesting recent gene flow. The coupling of numerous independent markers with extensive geographic sampling and simulations utilized here provides a framework for assessing the prevalence of gene flow in recently diverged species.     

Genetic and morphological studies of Nothobranchius (Cyprinodontiformes) from Malawi with description of Nothobranchius wattersi sp. nov.

Molecular and morphological data were used to explore evolutionary differentiation among populations of Nothobranchius in the Lake Malawi–upper Shire River and the Lakes Chilwa–Chiuta drainage systems in Malawi. The aim of the study was to test the hypothesis that Nothobranchius of the Malawi–Shire system constitute a separate evolutionary group from Nothobranchius kirki. Mitochondrial and nuclear sequence data show a strongly supported phylogenetic split into two monophyletic groups separating the Lake Malawi basin fish from N. kirki. Unlike N. kirki, Lake Malawi–Shire fish do not deviate from neutrality and express an excess of rare haplotypes and mutations in terminal branches, characteristic of recently expanded populations. Further, the two groups significantly differ in morphology. Two body characters (dorsal‐fin base length and pre‐pelvic–pre‐anal distance) are significantly different between the two species in both sexes. Several other characters are significantly different in either male or female comparisons with respect to both standard and head lengths, and robust morphological differentiation is detected by multivariate analysis. The two groups are readily distinguished on the basis of male colouration, especially in scale centres and the caudal fin. On the basis of this differentiation at the molecular and morphological levels, in addition to colouration, the Lake Malawi–Shire fish are hereby formally recognized as constituting a new species, Nothobranchius wattersi. This distinction is in agreement with the geomorphologic and recent climatic history in the region.     

Establishment and expansion of Lake Malawi rock fish populations after a dramatic Late Pleistocene lake level rise

Major environmental events that fragment populations among multiple island habitats have potential to drive large-scale episodes of speciation and adaptive radiation. A recent palaeolimnological study of sediment cores indicated that Lake Malawi underwent major climate-driven desiccation events 75 000–135 000 years ago that lowered the water level to at least 580 m below the present state and severely reduced surface area. After this period, lake levels rose and stabilized, creating multiple discontinuous littoral rocky habitats. Here, we present evidence supporting the hypothesis that establishment and expansion of isolated philopatric rock cichlid populations occurred after this rise and stabilization of lake level. We studied the Pseudotropheus ( Maylandia ) species complex, a group with both allopatric and sympatric populations that differ in male nuptial colour traits and tend to mate assortatively. Using coalescent analyses based on mitochondrial DNA, we found evidence that populations throughout the lake started to expand and accumulate genetic diversity after the lake level rise. Moreover, most haplotypes were geographically restricted, and the greatest genetic similarities were typically among sympatric or neighbouring populations. This is indicative of limited dispersal and establishment of assortative mating among populations following the lake level rise. Together, this evidence is compatible with a single large-scale environmental event being central to evolution of spatial patterns of genetic and species diversity in P. ( Maylandia ) and perhaps other Lake Malawi rock cichlids. Equivalent climate-driven pulses of habitat formation and fragmentation may similarly have contributed to observed rapid and punctuated cladogenesis in other adaptive radiations.     

Genomics of invasion: diversity and selection in introduced populations of monkeyflowers (Mimulus guttatus)

Global trade and travel is irreversibly changing the distribution of species around the world. Because introduced species experience drastic demographic events during colonization and often face novel environmental challenges from their native range, introduced populations may undergo rapid evolutionary change. Genomic studies provide the opportunity to investigate the extent to which demographic, historical and selective processes shape the genomic structure of introduced populations by analysing the signature that these processes leave on genomic variation. Here, we use next‐generation sequencing to compare genome‐wide relationships and patterns of diversity in native and introduced populations of the yellow monkeyflower (Mimulus guttatus). Genome resequencing data from 10 introduced populations from the United Kingdom (UK) and 12 native M. guttatus populations in North America (NA) demonstrated reduced neutral genetic diversity in the introduced range and showed that UK populations are derived from a geographic region around the North Pacific. A selective‐sweep analysis revealed site frequency changes consistent with selection on five of 14 chromosomes, with genes in these regions showing reduced silent site diversity. While the target of selection is unknown, genes associated with flowering time and biotic and abiotic stresses were located within the swept regions. The future identification of the specific source of origin of introduced UK populations will help determining whether the observed selective sweeps can be traced to unsampled native populations or occurred since dispersal across the Atlantic. Our study demonstrates the general potential of genome‐wide analyses to uncover a range of evolutionary processes affecting invasive populations.     

Variation in body size and trophic morphology within and among genetically differentiated populations of the cichlid fish, Metriaclima zebra, from Lake Malawi

1. The cichlid fish Metriaclima zebra , common in Lake Malawi, feeds by filtering plankton from the water and by brushing items from sediment covered substrata. It inhabits isolated rocky reefs among which community structure, resource availability and gene pools are likely to differ. We speculated that body size and trophic morphology of M. zebra might vary concomitantly.
2. We quantified the extent of genetic, body size and trophic variation within and between populations of M. zebra from southern Lake Malawi. Specifically, we tested the hypotheses that: (i) local populations are genetically differentiated, (ii) local populations differ in jaw morphology, dentition and standard length (SL), and (iii) variation in size is correlated with variation in trophic morphology.
3. Local populations of M. zebra differed in mean SL and were genetically differentiated. Moreover, populations exhibited dissimilar oral jaw morphologies and dentitions, perhaps related to differences in feeding biology. Variation in jaw shape was largely restricted to the curvature of the distal tip of the dentary. Populations were characterised by individuals with oblique, upward or downward directed gapes. Dental patterns differed in the proportion of unicuspid teeth in all rows of each jaw (dentaries and premaxillae) and the spacing of teeth in affected rows.
4. Within populations, jaw and tooth shapes were correlated with body size. Smaller individuals possessed upward curving jaws and closely packed multicusped teeth, while larger individuals exhibited relatively downward-directed jaws with increasing numbers of widely spaced unicuspid teeth.
5. Metriaclima zebra populations have increased in mean SL over the last decade, in contrast to a decline among Lake Malawi pelagic cichlids. Differences in size may contribute to variation in trophic morphology and may track local environmental dynamics in this lacustrine system.     

Genetically monomorphic invasive populations of the rapa whelk,Rapana venosa

Rapana venosa is a predatory marine gastropod native to the coastal waters of China, Korea, and Japan. Since the 1940s, R. venosa has been transported around the globe and introduced populations now exist in the Black Sea, the Mediterranean Sea, the Adriatic and Aegean seas, off the coasts of France and the Netherlands, in Chesapeake Bay, Virginia, USA, and in the Rio de la Plata between Uruguay and Argentina. This study surveyed variation in two mitochondrial gene regions to investigate the invasion pathways of R. venosa, identify likely sources for introduced populations, and evaluate current hypotheses of potential transportation vectors. Sequence data were obtained for the mitochondrial cytochrome c oxidase I and NADH dehydrogenase subunit 2 gene regions of 178 individuals from eight native locations and 106 individuals from 12 introduced locations. Collections from within the native range displayed very high levels of genetic variation while collections from all introduced populations showed a complete lack of genetic diversity; a single haplotype was common to all introduced individuals. This finding is consistent with the hypothesis that R. venosa was initially introduced into the Black Sea, and this Black Sea population then served as a source for the other secondary invasions by various introduction vectors including ballast water transport. Although non‐native R. venosa populations currently appear to be thriving in their new environments, the lack of genetic variability raises questions regarding the evolutionary persistence of these populations.     

Can chaoborid larvae control their zooplanktonic prey populations in a neotropical lake?

Although it is well known that predatory Chaoborus Lichtenstein 1800 larvae have a strong negative impact on zooplankton populations in many Holarctic lakes, this top‐down control is still not entirely clear in the Neotropics. In Lake Carioca, a Brazilian monomictic lake, chaoborids became one of the main zooplankton predators after the depletion of planktivorous fish by non‐native piscivorous fish. The aim of this research was to test the hypothesis that Chaoborus spp. larvae control the density of their prey populations. Enclosures simulated conditions with higher (Pr+) and lower (Pr?) predatory Chaoborus spp. larva densities over 30 days in two periods (stratification and mixing). Our results indicated that the dominant copepod species was affected only in the stratification period when its population was smaller. In this circumstance, predated individuals were probably not replaced in the same proportion by reproductive output of the smaller population. Rotifers were not sensitive to changes in chaoborids densities in both periods. The lack of a strong top‐down effect recorded by our mesocosm experiments emphasises the importance of continuing to investigate the ecological roles played by this invertebrate predator in aquatic food webs and if its ability to regulate the zooplankton in most Holarctic lakes can (or cannot) be widely generalised to Neotropical communities. Such a result also suggests that alternative research approaches, besides in‐situ experiments, may be necessary to better clarify this topic. Specifically for cladocerans, parallel paleolimnological analyses in Lake Carioca have supported the idea of a possible negative impact of Chaoborus larvae on bosminids after the introduction of non‐native piscivorous fishes. Abstract in Portuguese is available with online material.     

Introductions and extinction of fish in the African great lakes

The great lakes of Africa contain one of the most remarkable known examples of rapid evolution and speciation of a vertebrate group. The three major lakes - Lake Victoria, Lake Malawi and Lake Tanganyika - each contain a unique radiation of fish belonging to the family Cichlidae. This has produced species 'flocks' that are unique to each lake in species numbers and diversity. At present most of the cichlid species in Lake Victoria are facing extinction as a result of the introduction of an exotic fish species. There are many other examples of introductions or invasions of non-native fish that have had severe effects on the native fish populations and regional economics. These examples should provide a clear warning against the proposed introductions of exotic fish into Lake Malawi.     

Growth and reproduction of the non‐native icefish Neosalanx taihuensis Chen, 1956 (Salangidae) in a plateau lake,southwestern China

Growth, reproduction and abundance traits of the invasive icefish Neosalanx taihuensis Chen, 1956 were investigated monthly from July 2009 to May 2011 in Lake Erhai on the Yunnan‐Guizhou Plateau, south‐western China, in order to explore the changes in life‐history traits after translocation. The results indicated that the icefish exhibited obvious plasticity in growth and reproduction traits. Growth of the fish in Lake Erhai was faster than that in native waters and in other translocated reservoirs. By fitting the von Bertalanffy growth model to the data, it was estimated that icefish obtain an asymptotic size of 96.12 mm, a K of 1.61, and a t₀ of ‐0.26; the calculated overall growth performance index φ′ was 4.17. The strategy of reproduction changed from multiple‐ to single‐spawning. The spawning period was from October to December with the absolute and relative fecundities of 1250 ± 169 eggs per ind and 2557 ± 245 eggs per g, respectively. Plasticity in icefish growth and reproduction in Lake Erhai greatly facilitated its population establishment, making it one of the most abundant fish species. The icefish invasion in the lake may be one of the reasons for the decrease or extinction of native fish species populations, and some measures for the control of this invasive fish are suggested.     

Genetic structure of a native cyprinid in a reservoir‐altered stream network

1. Reservoirs modify riverine ecosystems worldwide, and often with deleterious impacts on native biota. The immediate effects of reservoirs on native fish species below dams and in impounded reaches have received considerable attention, but it is unclear how reservoirs may affect fish species at larger spatial and temporal scales. Documented declines of stream fish populations in direct tributaries of reservoirs suggest reservoir pools may reduce gene flow among historically connected populations. 2. Because of increased predator densities in reservoirs and the extent of habitat alteration in impounded reaches, I predicted reservoir habitats would reduce gene flow among small‐bodied fish populations separated by reservoir habitat. I used microsatellite markers to assess the spatial genetic structure of populations of the red shiner (Cyprinella lutrensis), in a reservoir‐fragmented stream network (Lake Texoma, U.S.A.). I also tested the prediction that populations in two direct tributaries that have experienced population declines would have low genetic diversity. Individuals were collected from six sites upstream of the reservoir, three sites in the reservoir and three sites in direct tributaries of the reservoir during 2008 and 2009. 3. Results indicate that most populations were isolated by distance with little divergence among populations. In one direct tributary population, however, there was substantial genetic divergence, and genetic diversity was significantly lower than in other populations. Gene flow also seemed to be lower in reservoir habitats than in intact stream habitats, suggesting reservoir habitats may be reducing gene flow among the reservoir‐separated populations. These results indicate that reservoirs may reduce gene flow among reservoir‐fragmented stream fish populations, altering the evolutionary trajectories of fragmented populations.     

Migratory behaviour shapes spatial genetic structure of cyprinid fishes within the Lake Malawi catchment

• Genetic differences among freshwater fish populations are dependent on life‐history characteristics of the species, including the range of adult dispersal and the extent of homing to natal breeding grounds. However, the effects of variation in such characteristics on population genetic connectivity are rarely studied comparatively among closely related species.
• We studied population genetic structure within three congeneric cyprinid species from the Lake Malawi catchment that differ substantially in life‐history traits and conservation status, using a combination of microsatellite and mitochondrial DNA markers. Mpasa (Opsaridium microlepis) is a large (70 cm total length) migratory species that spawns in rivers, but as an adult is exclusively known from the main lake body. Sanjika (Opsaridium microcephalum), is a medium size (30 cm total length) species that exists in lake breeding, river‐lake migratory and apparently landlocked populations. Dwarf sanjika (Opsaridium tweddleorum) is a small non‐migratory species (15 cm total length) that persists in small tributaries surrounding the main lake and adjoining rivers.
• The results revealed striking differences among the three species in spatial genetic structuring. The river‐lake migratory mpasa showed only weak yet significant population genetic structure within the main Lake Malawi catchment, suggesting that there is no strong natal homing. The habitat‐generalist sanjika showed only weak spatial genetic differentiation at microsatellite loci within the Lake Malawi catchment, but moderate structure in mitochondrial DNA, potentially reflecting male‐biased dispersal. The river‐restricted dwarf sanjika showed strong genetic structure in both microsatellite and mitochondrial DNA, suggesting strictly limited dispersal at both adult and juvenile stages.
• We conclude that contrasting migration life histories have resulted in dramatically different patterns of population genetic structure among these congeneric species. The observed patterns demonstrate how divergent life‐history evolution may strongly influence broader patterns of population genetic connectivity in freshwater fish, with consequences for management and conservation. Specifically the results suggesting gene flow among Lake Malawi populations of mpasa, an IUCN red‐listed ‘Endangered’ species endemic to the lake catchment, imply that conservation initiatives operating at both local and catchment scales are needed to reverse local population decline.

     

Genetic differentiation in Hippocrepis emerus (Leguminosae): allozyme and DNA fingerprint variation in disjunct Scandinavian populations

The structure of genetic variation in disjunct Scandinavian populations of Hippocrepis emerus was studied using allozymes and DNA fingerprinting. Variation in the three native regional populations in Scandinavia was compared with that in a recently introduced population in Sweden. In contrast to the recently introduced population, the native Scandinavian isolates of H. emerus showed high levels of allozyme fixation and low levels of DNA diversity. Variation in allozymes and at DNA fingerprint loci showed closely congruent patterns of geographic variation, with pronounced differentiation between the native Norwegian and Swedish isolates of the species. The structure of genetic variation in native Scandinavian H. emerus is interpreted in terms of historical population bottlenecks and founder events during the species' postglacial immigration into Scandinavia.     

Genetic structure and gene flow in an endangered native tilapia fish (Oreochromis esculentus) compared to invasive Nile tilapia (Oreochromis niloticus) in Yala swamp, East Africa

The introduction of invasive Nile tilapia (Oreochromis niloticus), and the rapacious predator Nile perch (Lates niloticus), into Lake Victoria resulted in a decline in population sizes, genetic diversity and even extirpation of native species which were previously the mainstay of local fisheries. However, remnant populations of native fish species, including tilapia, still persist in satellite lakes around Lake Victoria where they may coexist with O. niloticus. In this study we assessed population genetic structure, diversity, and integrity of the native critically endangered Singidia tilapia (O. esculentus) in its refugial populations in the Yala swamp, Kenya, and contrasted this diversity with populations of the invasive tilapia O. niloticus in satellite lakes (Kanyaboli, Namboyo and Sare) and Lake Victoria. Based on mtDNA control region sequences and eight nuclear microsatellite loci, we did not detect any mtDNA introgression between the native and the invasive species in Lakes Kanyaboli and Namboyo, but did find low levels of nuclear admixture, primarily from O. niloticus to O. esculentus. Some genetic signal of O. esculentus in O. niloticus was found in Lake Sare, where O. esculentus is not found, suggesting it has recently been extirpated by the O. niloticus invasion. In both species, populations in the satellite lakes are significantly genetically isolated from each other, with private mtDNA haplotypes and microsatellite alleles. For O. niloticus, genetic diversity in satellite lakes was similar to that found in Lake Victoria. Our data imply a low frequency of immigration exchange between the two populations of O. esculentus and we suggest that the populations of this endangered species and important fisheries resource should be conserved separately in Lakes Kanyaboli and Namboyo and with high priority.     

Resistance of an invasive gastropod to an indigenous trematode parasite in Lake Malawi

Successful establishment and spread of biological invaders may be promoted by the absence of population-regulating enemies such as pathogens, parasites or predators. This may come about when introduced taxa are missing enemies from their native habitats, or through immunity to enemies within invaded habitats. Here we provide field evidence that trematode parasites are absent in a highly invasive morph of the gastropod Melanoides tuberculata in Lake Malawi, and that the invasive morph is resistant to indigenous trematodes that castrate and induce gigantism in native M. tuberculata. Since helminth infections can strongly influence host population abundances in other host-parasite systems, this enemy release may have provided an advantage to the invasive morph in terms of reproductive capacity and survivorship.