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Parasites may have strong eco-evolutionary interactions with their hosts. Consequently, they may contribute to host diversification. The radiation of cichlid fish in Lake Victoria provides a good model to study the role of parasites in the early stages of speciation. We investigated patterns of macroparasite infection in a community of 17 sympatric cichlids from a recent radiation and 2 older species from 2 nonradiating lineages, to explore the opportunity for parasite-mediated speciation. Host species had different parasite infection profiles, which were only partially explained by ecological factors (diet, water depth). This may indicate that differences in infection are not simply the result of differences in exposure, but that hosts evolved species-specific resistance, consistent with parasite-mediated divergent selection. Infection was similar between sampling years, indicating that the direction of parasite-mediated selection is stable through time. We morphologically identified 6 Cichlidogyrus species, a gill parasite that is considered a good candidate for driving parasite-mediated speciation, because it is host species-specific and has radiated elsewhere in Africa. Species composition of Cichlidogyrus infection was similar among the most closely related host species (members of the Lake Victoria radiation), but two more distantly related species (belonging to nonradiating sister lineages) showed distinct infection profiles. This is inconsistent with a role for Cichlidogyrus in the early stages of divergence. To conclude, we find significant interspecific variation in parasite infection profiles, which is temporally consistent. We found no evidence that Cichlidogyrus-mediated selection contributes to the early stages of speciation. Instead, our findings indicate that species differences in infection accumulate after speciation.

Species interactions are a key aspect of evolutionary biology. Parasites, specifically, are drivers of the evolution of species communities and impact biosecurity and public health. However, when using interaction networks for evolutionary studies, interdependencies between distantly related species in these networks are shaped by ancient and complex processes. We propose using recent interacting host-parasite radiations, e.g. African cichlid fishes and cichlid gill parasites belonging toCichlidogyrus(Dactylogyridae, Monogenea), as macroevolutionary model of species interactions. The cichlid-Cichlidogyrusnetwork encompasses 138 parasite species and 416 interactions identified through morphological characteristics and genetic markers in 160 publications. We discuss the steps required to develop this model system based on data resolution, sampling bias, and reporting quality. In addition, we propose the following steps to guide efforts for a macroevolutionary model system for species interactions: first, evaluating and expanding model system outcome measures to increase data resolution; second, closing knowledge gaps to address underreporting and sampling bias arising from limited human and financial resources. Identifying phylogenetic and geographic targets, creating systematic overviews, enhancing scientific collaborations, and avoiding data loss through awareness of predatory journal publications can accelerate this process; and third, standardising data reporting to increase reporting quality and to facilitate data accessibility.

The Lower Congo Basin is characterized by a mangrove-lined estuary at its mouth and, further upstream, by many hydrogeographical barriers such as rapids and narrow gorges. Five localities in the mangroves and four from (upstream) left bank tributaries or pools were sampled. On the gills of Coptodon tholloni, Coptodon rendalli, Hemichromis elongatus, Hemichromis stellifer and Tylochromis praecox, 17 species of parasites (Dactylogyridae & Gyrodactylidae, Monogenea) were found, eight of which are new to science. Six of these are herein described: Cichlidogyrus bixlerzavalai n. sp. and Cichlidogyrus omari n. sp. from T praecox, Cichlidogyrus calycinus n. sp. and Cichlidogyrus polyenso n. sp. from H. elongatus, Cichlidogyrus kmentovae n. sp. from H. stellifer and Onchob-della ximenae n. sp. from both species of Hemichromis. On Cichlidogyrus reversati a ridge on the accessory piece was discovered that connects to the basal bulb of the penis. We report a putative spillback effect of the native parasites Cichlidogyrus berradae, Cichlidogyrus cubitus and Cichlidogyrus flexicolpos from C. tholloni to the introduced C. rendalli. From our results, we note that the parasite fauna of Lower Congo has a higher affinity with the fauna of West African and nearby freshwater ecoregions than it has with fauna of other regions of the Congo Basin and Central Africa.

Thanks to high species diversity and a broad range of speciation mechanisms, cichlid fishes represent a textbook model in evolutionary biology. They are also of substantial economic value. Despite this importance, cichlid parasites remain understudied, although some are more diverse than their hosts. They may offer important insights into cichlid evolution and the evolution of host-parasite interactions. We review five major lines of research conducted on cichlid parasites so far: the study of parasite diversity and speciation; the role of parasites in cichlid diversification; the evolutionary ecology of host specificity; historical biogeography; and biological invasions. We call for more research in these areas and suggest approaches to valorise the potential that cichlid parasites hold for the study of evolutionary parasitology.

Background: Parasite switches to new host species are of fundamental scientific interest and may be considered an important speciation mechanism. For numerous monogenean fish parasites, infecting different hosts is associated with morphological adaptations, in particular of the attachment organ (haptor). However, haptoral morphology in Cichlidogyrus spp. (Monogenea, Dactylogyridea), parasites of African cichlids, has been mainly linked to phylogenetic rather than to host constraints. Here we determined the position of Cichlidogyrus amieti, a parasite of species of Aphyosemion (Cyprinodontiformes, Nothobranchiidae) in the phylogeny of its congeners in order to infer its origin and assess the morphological changes associated with host-switching events. Methods: The DNA of specimens of C. amieti isolated from Aphyosemion cameronense in Cameroon was sequenced and analyzed together with that of Cichlidogyrus spp. from cichlid hosts. In order to highlight the influence of the lateral transfer of C. amieti on the haptoral sclerotised parts we performed a Principal Component Analysis (PCA) to compare the attachment organ structure of C. amieti to that of congeners infecting cichlids. Results: Cichlidogyrus amieti was found to be nested within a strongly supported clade of species described from Hemichromis spp. (i.e. C. longicirrus and C. dracolemma). This clade is located at a derived position of the tree, suggesting that C. amieti transferred from cichlids to Cyprinodontiformes and not inversely. The morphological similarity between features of their copulatory organs suggested that C. amieti shares a recent ancestor with C. dracolemma. It also indicates that in this case, these organs do not seem subjected to strong divergent selection pressure. On the other hand, there are substantial differences in haptoral morphology between C. amieti and all of its closely related congeners described from Hemichromis spp.. Conclusions: Our study provides new evidence supporting the hypothesis of the adaptive nature of haptor morphology. It demonstrates this adaptive component for the first time within Cichlidogyrus, the attachment organs of which were usually considered to be mainly phylogenetically constrained.

A major goal in ecology is to unravel how species assemblages emerge and how they are structured across the landscape. Host-parasite systems are particularly interesting in this context, as limited host dispersal may promote the differentiation of parasite communities. We examined whether the patterns of species diversity in Cichlidogyrus, a genus of monogenean parasitic flatworms with a direct life cycle, are consistent with the hypothesis that parasite diversity is driven by host dispersal. This was carried out by comparing two sympatric cichlid hosts (Tropheus moorii and Simochromis diagramma) with contrasting dispersal abilities. Genetic connectivity among host populations along the Zambian shoreline of Lake Tanganyika was estimated using microsatellite genotyping. Cichlidogyrus parasites were isolated and identified morphologically to the species level.Simochromis diagramma, a host with a high dispersal capacity, was infected by a low number of Cichlidogyrus species, and the parasite assemblages were similar among host populations. In contrast, T.moorii, a host with a low dispersal capacity, was infected by a large number of Cichlidogyrus species, and the parasite assemblages differed strongly among host populations. These outcomes were thus as expected from the hypothesis. Because of the strong host specificity of these Cichlidogyrus species, a lack of connectivity among host populations might facilitate allopatric speciation of the parasite.

Background: Adaptation to different ecological environments is thought to drive ecological speciation. This phenomenon culminates in the radiations of cichlid fishes in the African Great Lakes. Multiple characteristic traits of cichlids, targeted by natural or sexual selection, are considered among the driving factors of these radiations. Parasites and pathogens have been suggested to initiate or accelerate speciation by triggering both natural and sexual selection. Three prerequisites for parasite-driven speciation can be inferred from ecological speciation theory. The first prerequisite is that different populations experience divergent infection levels. The second prerequisite is that these infection levels cause divergent selection and facilitate adaptive divergence. The third prerequisite is that parasite-driven adaptive divergence facilitates the evolution of reproductive isolation. Here we investigate the first and the second prerequisite in allopatric chromatically differentiated lineages of the rock-dwelling cichlid Tropheus spp. from southern Lake Tanganyika (Central Africa). Macroparasite communities were screened in eight populations belonging to five different colour morphs. Results: Parasite communities were mainly composed of acanthocephalans, nematodes, monogeneans, copepods, branchiurans, and digeneans. In two consecutive years (2011 and 2012), we observed significant variation across populations for infection with acanthocephalans, nematodes, monogeneans of the genera Gyrodactylus and Cichlidogyrus, and the copepod Ergasilus spp. Overall, parasite community composition differed significantly between populations of different colour morphs. Differences in parasite community composition were stable in time. The genetic structure of Tropheus populations was strong and showed a significant isolation-by-distance pattern, confirming that spatial isolation is limiting host dispersal. Correlations between parasite community composition and Tropheus genetic differentiation were not significant, suggesting that host dispersal does not influence parasite community diversification. Conclusions: Subject to alternating episodes of isolation and secondary contact because of lake level fluctuations, Tropheus colour morphs are believed to accumulate and maintain genetic differentiation through a combination of vicariance, philopatric behaviour and mate discrimination. Provided that the observed contrasts in parasitism facilitate adaptive divergence among populations in allopatry (which is the current situation), and promote the evolution of reproductive isolation during episodes of sympatry, parasites might facilitate speciation in this genus.

Lake Tanganyika is the deepest and oldest African Great Lake and of economic importance. While the diversity of its endemic cichlid radiations yielded scientific interest, a number of cichlid tribes have few representatives in the lake. Some of those, namely Oreochromini (ex-Tilapiini), Haplochromini and Tylochromini, reach higher species numbers in riverine systems. Conversely, the phylogenetic position of the monospecific and endemic Boulengerochromini is unclear. The oreochromines Oreochromis tanganicae and Oreochromis niloticus, the haplochromine Astatotilapia burtoni, the tylochromine Tylochromis polylepis and the boulengerochromine Boulengerochromis microlepis, the largest cichlid species worldwide, were surveyed for ancyrocephalid monogenean gill parasites. Five new species are proposed. Cichlidogyrus gillardinae sp. n. is described from A. burtoni, Cichlidogyrus mbirizei sp. n. from O. tanganicae and Cichlidogyrus nshomboi sp. n. from B. microlepis. T. polylepis harbours Cichlidogyrus mulimbwai sp. n., Cichlidogyrus muzumanii sp. n. and a third, presently undescribed species. Four species known from outside the Tanganyika Basin were retrieved on the oreochromines. The host species are scientific models or important in the sectors of fisheries or ornamental fish trade. Moreover, their phylogenetic positions render them well-suited to help elucidate the historic relationships between riverine and lacustrine African cichlids. In this framework, their Cichlidogyrus fauna is compared to congeners known from African rivers and to the few Tanganyika representatives described. While the parasites of Oreochromis, A. burtoni and T. polylepis are reminiscent of those infecting related hosts throughout Africa, B. microlepis hosts a Cichlidogyrus morphotype typical of Lake Tanganyika. This supports its placement within an endemic cichlid radiation.

Examination of gill parasites from Ophthalmotilapia ventralis (Boulenger, 1898) in Lake Tanganyika (Africa) revealed the presence of four new species of Monogenea, all members of Cichlidogyrus Paperna, 1960 (Ancyrocephalidae). In view of the systematic importance of haptoral structure, the fish host shows a remarkable diversity of morphological groups. Cichlidogyrus vandekerkhovei sp. nov. and C. makasai sp. nov. are especially characterized by the unusual length of the dorsal transverse bar auricles, while C. sturmbaueri sp. nov. is distinguished by the unique shape of the accessory piece of its male copulatory organ. Importantly, C. centesimus sp. nov. displays a number of features new to the genus, namely a spirally coiled thickening at the end of the penis, the absence of an accessory piece in the genital apparatus, and a hitherto unknown uncinuli configuration in the haptor. This is the first record of ancyrocephalid parasites from the Tanganyika basin. Some mechanisms possibly contributing to this yet unknown diversity are discussed, identifying topics deserving further scientific scrutiny.