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Citation: Mushagalusa Mulega, A.; Van Steenberge, M.; Kmentová, N.; Muterezi Bukinga, F.; Rahmouni, I.; Masilya, P.M.; Benhoussa, A.; Pariselle, A.; Vanhove, M.P.M. Abstract: The ichthyofauna of Lake Tanganyika consists of 12 families of fish of which five belong to Siluriformes (catfishes). Studies on Siluriformes and their parasites in this lake are very fragmentary. The present study was carried out to help fill the knowledge gap on the monogeneans infesting the siluriform fishes of Lake Tanganyika in general and, more particularly, Clarias gariepinus. Samples of gills of Clarias gariepinus (Clariidae) were examined for ectoparasites. We identified the monogenean Gyrodactylus transvaalensis (Gyrodactylidae). This is the first time this parasite was found infecting gills. We are the first to observe a large spine in the male copulatory organ of this species and to provide measurements of its genital spines; this completes the description of the male copulatory organ, which is important in standard monogenean identification. This is the first monogenean species reported in C. gariepinus at Lake Tanganyika and the third known species on a representative of Siluriformes of this lake. It brings the total number of species of Gyrodactylus recorded in Lake Tanganyika to four. Knowing that other locations where this species has been reported are geographically remote from Lake Tanganyika, we propose a "failure to diverge" phenomenon for G. transvaalensis.

Lake Tanganyika, East Africa, is the oldest and deepest African Great Lake and harbours one of the most diverse fish assemblages on earth. Two clupeid fishes, Limnothrissa miodon and Stolothrissa tanganicae, constitute a major part of the total fish catch, making them indispensable for local food security. Parasites have been proposed as indicators of stock structure in highly mobile pelagic hosts. We examined the monogeneans Kapentagyrus limnotrissae and Kapentagyrus tanganicanus (Dactylogyridae) infecting these clupeids to explore the parasites' lake-wide population structure and patterns of demographic history. Samples were collected at seven sites distributed across three sub-basins of the lake. Intraspecific morphological variation of the monogeneans (n = 380) was analysed using morphometrics and geomorphometrics of sclerotised structures. Genetic population structure of both parasite species (n = 246) was assessed based on a 415 bp fragment of the mitochondrial cytochrome c oxidase subunit I (COI) gene. Overall, we observed a lack of clear geographical morphological differentiation in both parasites along a north-south axis. This lack of geographical population structure was also reflected by a large proportion of shared haplotypes, and a pattern of seemingly unrestricted gene flow between populations. Significant morphological and genetic differentiation between some populations might reflect temporal differentiation rather than geographical isolation. Overall, the shallow population structure of both species of Kapentagyrus reflects the near-panmictic population structure of both host species as previously reported. Morphological differences related to host species identity of K. tanganicanus were consistent with incipient speciation at the genetic level. Both parasite species experienced a recent demographic expansion, which might be linked to paleohydrological events. Finally, interspecific hybridisation was found in Kapentagyrus, representing the first case in dactylogyrid monogeneans. (C) 2020 Australian Society for Parasitology. Published by Elsevier Ltd. All rights reserved. The authors would like to thank L. Raisingerová, C. Rahmouni, H. Zimmermann, A. P. H. Bose and W. Salzburger for help with collecting host samples. M. Jorissen is gratefully acknowledged for fruitful discussions about monogenean morphological variability and help in the laboratory. Special thanks to the staff of the parasitological group at Masaryk University, Brno (Czech Republic), the Research Group Zoology at Hasselt University (Belgium), Research Centre of Hydrobiology in Uvira (DR Congo) and Fisheries Research Unit in Mpulungu (Zambia) for their hospitality. Research was supported by the Czech Science Foundation (P505/12/G112 (ECIP) and GA19-13573S), EMBRC Belgium – FWO project GOH3817N as well as by a joint program between the Austrian Agency for International Cooperation in Education and Research (OEAD; project number CZ 08/2018; to SK) and the Ministry of Education, Youth and Sports (project number 8J18AT007). MPMV was co-financed by institutional funding of the Finnish Museum of Natural History and the Belgian Directorate-General for Development Cooperation and Humanitarian Aid (CEBioS program). ELRDK was supported by the Belgian Development Cooperation through VLIR-UOS (VLADOC scholarship NDOC2016PR006 to ELRDK and South Initiative project CD2018SIN218A101). Fieldwork was carried out with the approval of the competent local authorities under mission statement 031/MINRST/CRH-U/2016 and the permission of the Fisheries Department of Zambia and under a study permit issued by the government of Zambia (SP 008732).

In the framework of the study of Siluriform fish monogeneans of Lake Tanganyika, we described two new species of Bagrobdella from Auchenoglanis occidentalis, Bagrobdella vanhovei sp. nov. is characterized by the morphology of its MCO which is unique (terminal opening) and Bagrobdella vansteenbergei sp. nov. characterized by the size of its hooks, which are almost all of the same size, and its male copulating organ with a unique shape: sub-terminal opening and no membrane surrounding.

Metazoan parasites encompass a significant portion of the global biodiversity. Their relevance for environmental and human health calls for a better understanding as parasite macroevolution remains mostly understudied. Yet limited molecular, phenotypic, and ecological data have so far discouraged complex analyses of evolutionary mechanisms and encouraged the use of data discretisation and body-size correction. In this case study, we aim to highlight the limitations of these methods and propose new methods optimised for small datasets. We apply multivariate phylogenetic comparative methods (PCMs) and statistical classification using support vector machines (SVMs) to a data-deficient host-parasite system. We use continuous morphometric and host range data currently widely inferred from a species-rich lineage of parasites (Cichlidogyrus incl. Scutogyrus - Platyhelminthes: Monogenea, Dactylogyridae) infecting cichlid fishes. For PCMs, we modelled the attachment organ and host range evolution using the data of 135 species and an updated multi-marker (28S and 18S rDNA, ITS1, COI mtDNA) phylogenetic reconstruction of 58/137 described species. Through a cluster analysis, SVM-based classification, and taxonomic literature survey, we infered the systematic informativeness of discretised and continuous characters. We demonstrate that an update to character coding and size-correction techniques is required as some techniques mask phylogenetic signals but remain useful for characterising species groups of Cichlidogyrus. Regarding the attachment organ evolution, PCMs suggest a pattern associated with genetic drift. Yet host and environmental parameters might put this structure under stabilising selection as indicated by a limited morphological variation. This contradiction, the absence of a phylogenetic signal and multicollinearity in most measurements, a moderate 73% accordance rate of taxonomic approach and SVMs, and a low phylogenetic informativeness of reproductive organ data suggest an overall limited systematic value of the measurements included in most species characterisations. We conclude that PCMs and SVM-based approaches are suitable tools to investigate the character evolution of data-deficient taxa.

BackgroundMonogenean parasites have never been formally reported on fish from the Lufira basin. Then it is hypothesised that multiple monogenean species are to be recorded that are new to the region. This study aimed to record the gill monogenean parasite fauna of three cichlid fish species in the Upper Lufira basin by inventorying their diversity (species composition) and analysing their infection parameters (prevalence, mean intensity and abundance).MethodsOreochromis mweruensis, Coptodon rendalli, and Serranochromis macrocephalus were selected for the study, given their economic value and their abundance in the Upper Lufira basin. Monogeneans were isolated from the gills and stomach, mounted on glass slides with either Hoyer’s medium or ammonium picrate-glycerin for further identification under a stereomicroscope, based on morphological analysis of genital and haptoral hard parts. Indices of diversity and infections parameters were calculated.ResultsA total of thirteen gill monogenean parasite species (Cichlidogyrus dossoui, C. halli, C. karibae, C. mbirizei, C. papernastrema, C. quaestio, C. sclerosus, C. tiberianus, C. tilapiae, C. zambezensis, Scutogyrus gravivaginus, S. cf. bailloni and Gyrodactylus nyanzae) and one stomach monogenean (Enterogyrus malmbergi) were reported. A species richness of S= 10 for O. mweruensis, S= 6 for C. rendalli and S= 2 for S. macrocephalus were recorded. Five parasite species were reported to be common amongst O. mweruensis and C. rendalli. The most prevalent parasite species were C. halli (P= 80.9%) on O. mweruensis, C. dossoui (P= 92.9%) on C. rendalli and C. karibae and C. zambezensis (both of which P = 9.1%) on S. macrocephalus with a respective mean infection intensity of 7.9 on O. mweruensis, 9.8 on C. rendalli and 5 and 15, respectively, on S. macrocephalus. Results of this study reported new host ranges for five parasites species (C. quaestio, S. cf. bailloni, E. malmbergi on O. mweruensis, C. halli on C. rendalli and C. karibae on S. macrocephalus) as well as new geographical records for three of them (S. cf. bailloni, E. malmbergi, C. karibae).ConclusionsThis study highlights the richness of monogenean communities in the Upper Lufira basin and is a starting point for future helminthological studies, e.g. on the use of fish parasites as indicators of anthropogenic impacts.

In the framework of the study of Siluriform fish monogeneans of Lake Tanganyika, we described two new species of Bagrobdella Paperna, 1969 from Auchenoglanis occidentalis (Valenciennes, 1840). Bagrobdella vanhovei sp. nov. is characterized by the morphology of its MCO which is unique among its congeners, presenting a non-terminal opening, whereas the other species have a terminal opening, and Bagrobdella vansteenbergei sp. nov. characterized by the size of its hooks, which are almost all of the same size, and its male copulating organ with a unique shape: a sub-terminal opening and no membrane surrounding. The Multivariate analysis done on morphometrical characters shows that the new and already described species are well individualized, except for Bagrobdella parauchenoglanii Akoumba, Pariselle, Tombi & Bilong Bilong, 2017 and Bagrobdella fraudulenta Euzet & Le Brun, 1990 (but these two species are easily distinguishable by their morphology), and that B. vanhovei sp. nov. has a great intra-specific morphometrical variation.

Nile tilapia, one of the most popular aquaculture species worldwide, has been introduced into the Congo Basin several times. In previous morphological studies, we showed that some of the monogenean gill parasites were co-introduced with Nile tilapia and some spilled over to native Congolese cichlids. In this study, we investigated the co-introduced monogeneans of Nile tilapia genetically from three major parts of the Congo Basin; Upper, Middle and Lower Congo. We generated sequences of Congolese native and introduced monogeneans from native and introduced tilapias and evaluate their position in a phylogeny. Additionally, we generated sequences of the same species of monogeneans co-introduced with Nile tilapia in Madagascar and of a native population of Nile tilapia from Burundi. Our results confirm the co-introductions in the Congo. We found that co-introduced parasites are less genetically diverse than native ones, and that there was no geographical pattern between introduced populations. Furthermore, our COI haplotype networks suggest multiple introduction events of Nile tilapia into the Congo Basin. Additionally, we tested the barcoding gap and the performance of mitochondrial COI and nuclear ribosomal ITS-1, 28S and 18S markers. We found a significant intra/interspecific barcoding gap of 15% for COI, but none for the other markers. Our molecular results reveal that Cichlidogyrus halli, C. papernastrema, C. tiberianus, C. cirratus and C. zambezensis are in need of taxonomic revision.

The fisheries of Lake Tanganyika play an important role in food security in Central and Eastern Africa. Conservation of these valuable fish stocks will benefit from documenting the ideas, opinions and observations of stakeholders. Knowledge of the perceptions and an understanding of the concerns and struggles of stakeholders of these fisheries can provide policy-makers with recommendations for more suitable fisheries management. We did 1018 interviews with stakeholders, in one close-ended and three semi-open ended surveys. Factor analysis revealed seven clusters of opinions. Linear-mixed effects models identified common grounds and differences in opinions between groups of stakeholders about strategies in fisheries management. Stakeholders of the fisheries indicated challenges due to weather or climate variability, a noticeable decrease in fish abundance and size, and an increase in the price of fish. Fishermen experienced a lack of safety on the lake, including aggression and dangerous weather conditions, and hardly had access to safety gear and infrastructure. Landing site officials, state employees who monitor the beaches, mentioned capture of juveniles and declining catch-rates as the biggest threats to the fisheries. None of the groups of stakeholders attributed the problems in the fisheries to overfishing or overpopulation. We found similarities in opinions over a wide range of stakeholder groups, with many stakeholders asking for better and fair enforcement of existing legislation. State employees were more positive than the other groups towards creating more strict regulation of the fisheries. The results presented offer focuspoints for policy-makers to improve the management of the Lake Tanganyika pelagic fisheries.

BackgroundMonogenea van Beneden, 1858 is a group of parasitic flatworms, commonly found infecting bony fish. Several genera, such asCichlidogyrusPaperna, 1960, are reported to include potential pathogenic species that can negatively impact aquaculture fish stocks. They can switch from introduced to native fish and vice versa. In Africa (and all over the world), fish species belonging to Cichlidae are often kept in aquaculture and represent a major source of food. Thus, research on the biodiversity and occurrence of monogenean species on these fish is of importance for aquaculture and conservation. The present study is a survey of the diversity of species ofCichlidogyrusin the south of the Democratic Republic of the Congo (DRC) on three cichlid species:Orthochromissp. ‘Lomami’,Serranochromiscf.macrocephalus, andTilapia sparrmaniiSmith, 1840.MethodsSpecimens ofCichlidogyruswere isolated from the gills and mounted on glass slides with Hoyer’s medium. The genital and haptoral hard parts were measured and drawn using interference contrast.ResultsIn total, six species ofCichlidogyruswere found, all new to science:C. bulbophallusn. sp. andC. pseudozambezensisn. sp. onS.cf.macrocephalus,C. flagellumn. sp. andC. lobusn. sp. onT. sparrmanii,C. ranulan. sp. onS.cf.macrocephalusandOrthochromissp. ‘Lomami’, andC. maeandern. sp. found onOrthochromissp. ‘Lomami’ andT. sparrmanii. The first four species are considered to be strict specialists,C. ranulan. sp. an intermediate generalist andC. maeandern. sp. a generalist. These parasite species show morphological similarities to species found in the Lower Guinea and Zambezi ichthyofaunal provinces, which might be explained by past river capture events between river systems of the Congo Province and both these regions.ConclusionsSerranochromiscf.macrocephalusandOrthochromissp. ‘Lomami’ can harbour respectively three and two species ofCichlidogyrus, all described in this study.Tilapia sparrmaniican harbour seven species, of which three are described in the present study. These results highlight the species diversity of this parasite genus in the Congo Basin.

Fish have been widely translocated into non-native areas, commonly as fishery targets. Since fish figure as hosts of various parasite taxa, their introduction may pose often-underestimated threats to ecosystems. However, parasites can also serve to track host species’ introduction routes when these would otherwise be unknown. To verify the potential of parasites in reconstructing invasion routes, we investigated two of the best-documented introductions: those of Limnothrissa miodon into lakes Kivu and Kariba. As a proof of concept, we investigate the possibility of using parasites to evaluate the effect of host size in the introduction pathway and to track the host origins of L. miodon. Combining historical collections and recent field samples, specimens of L. miodon from Lake Kivu and Lake Kariba were examined for monogenean flatworms. Intraspecific variation was investigated using morphometrics of the parasite’s sclerotised structures. Three markers from the ribosomal DNA region were used for genetic parasite identification. In Lake Tanganyika, L. miodon is infected by two species of monogeneans, Kapentagyrus limnotrissae and K. tanganicanus. One of these species, K. limnotrissae, was found on L. miodon from Lake Kariba. In contrast, not a single monogenean individual was found in specimens from Lake Kivu. Morphometric results suggested that the origin of K. limnotrissae introduced into Lake Kariba may be the southern part of Lake Tanganyika, which corresponds to historical reports. Moreover, differences in the size of introduced fish, fry versus juveniles, were proposed as one of the factors influencing parasite occurrence in non-native areas. This supports the potential use of monogeneans as markers for host origin. This study was supported by ECIP (European Centre of Ichthyoparasitology); project No. P505/12/G112. Work in the collections of RMCA (Royal Museum for Central Africa) was financed by the SYNTHESYS Project (http://www.synthesys.info/) (BE-TAF-5731) which is financed by European Community Research Infrastructure Action under the FP7 Integrating Activities Programme.

Whereas Lake Tanganyika’s littoral and benthic zones are famous for their diverse fish communities, its pelagic zone is dominated by few species, of which two representatives of Clupeidae (Limnothrissa miodon and Stolothrissa tanganicae) take a pivotal role. We investigated the monogenean fauna infecting these freshwater clupeids to explore the link between parasite morphology and host species identity, or seasonal and geographical origin, which may reveal host population structure. Furthermore, we conducted phylogenetic analyses to test whether these parasitic flatworms mirror their host species’ marine origin. Based on 406 parasite specimens infecting 385 host specimens, two monogenean species of Kapentagyrus Kmentová, Gelnar and Vanhove, gen. nov. were morphologically identified and placed in the phylogeny of Dactylogyridae using three molecular markers. One of the species, Kapentagyrus limnotrissae comb. nov., is host-specific to L. miodon while its congener, which is new to science and described as Kapentagyrus tanganicanus Kmentová, Gelnar and Vanhove, sp. nov., is infecting both clupeid species. Morphometrics of the parasites’ hard parts showed intra-specific variability, related to host species identity and seasonality in K. tanganicanus. Significant intra-specific differences in haptor morphometrics between the northern and southern end of Lake Tanganyika were found, and support the potential use of monogeneans as tags for host population structure. Based on phylogenetic inference, we suggest a freshwater origin of the currently known monogenean species infecting clupeids in Africa, with the two species from Lake Tanganyika representing a quite distinct lineage.

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 Onchobdella 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. Six espèces nouvelles de Dactylogyridae (Platyhelminthes, Monogenea) parasites des branchies de cichlidés (Teleostei, Cichliformes) du bassin du Bas Congo. Le cours du Bas Congo est caractérisé par un estuaire bordé de mangroves et, plus en amont, par de nombreuses barrières hydro-géographiques formées de rapides ou de gorges étroites. Nous avons échantillonné dans cinq localités au niveau des mangroves et dans quatre autres en amont sur la rive gauche du fleuve, dans des affluents ou des étangs. Nous avons trouvé, sur les branchies de Coptodon tholloni, Coptodon rendalli, Hemichromis elongatus, Hemichromis stellifer et Tylochromis praecox, 17 espèces de parasites (Dactylogyridae et Gyrodactylidae, Monogenea), dont huit sont nouvelles pour la science. Six d’entre elles sont décrites ici : Cichlidogyrus bixlerzavalai n. sp. et Cichlidogyrus omari n. sp. sur T. praecox, Cichlidogyrus calycinus n. sp. et Cichlidogyrus polyenso n. sp. sur H. elongatus, Cichlidogyrus kmentovae n. sp. sur H. stellifer et Onchobdella ximenae n. sp. sur les deux espèces d’Hemichromis. Nous décrivons, chez Cichlidogyrus reversati, une arête sur la pièce accessoire qui la connecte avec l’ampoule basale basal du pénis. Nous notons un probable transfert latéral des espèces autochtones Cichlidogyrus berradae, Cichlidogyrus cubitus et Cichlidogyrus flexicolpos de C. tholloni vers l’espèce introduite C. rendalli. Nos résultats montrent que la faune parasitaire du Bas Congo présente plus d’affinités avec les faunes d’Afrique de l’Ouest ou des écorégions d’eau douce voisines, qu’avec celles des autres parties du bassin du Congo ou de l’Afrique Centrale.

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.

Lake Tanganyika’s biodiversity and endemicity sparked considerable scientific interest. Its monogeneans, minute parasitic flatworms, have received renewed attention. Their host-specificity and simple life cycle render them ideal for parasite speciation research. Because of the wide ecological and phylogenetic range of its cichlids, Lake Tanganyika is a “natural experiment” to contrast factors influencing monogenean speciation. Three representatives of Bathybatini (Bathybates minor, B. fasciatus, B. vittatus), endemic predatory non-littoral cichlids, host a single dactylogyridean monogenean species. It is new to science and described as Cichlidogyrus casuarinus sp. nov. This species and C. nshomboi and C. centesimus, from which it differs by the distal end of the accessory piece of the male apparatus and the length of its heel, are the only Cichlidogyrus species with spirally coiled thickening of the penis wall. In Cichlidogyrus, this feature was only found in parasites of endemic Tanganyika tribes. The seemingly species-poor Cichlidogyrus community of Bathybatini may be attributed to meagre host isolation in open water. The new species infects cichlids that substantially differ phylogenetically and ecologically. This may be an adaptation to low host availability. Cichlidogyrus species infecting African Great Lake cichlids are summarized and proposed as model for the influence of host ecology on disease transmission.

Background. Despite the importance of Lake Tanganyika's biodiversity for science and the livelihoods of the riparian people, high-resolution surveys of the fish biodiversity are sparse and fragmentary, especially along the western (Congolese) shoreline. The coast suffers locally from intensive human activities and lacks adequate protective measures or nature reserves. However, in view of the intra-lacustrine endemism of this fish fauna, conservation needs to be managed lake-wide at a fine scale, necessitating detailed inventories on fish species distribution. The study aims at updating knowledge on fish diversity and distribution along the north-western and central western shores of Lake Tanganyika. Materials and methods. Fish specimens were collected using gill-and seine nets, by snorkelling and SCUBA diving, and through purchases on the local markets. Results. Over 28 locations were sampled, and 84 cichlid- and 30 non-cichlid fish species (belonging to Protopteridae, Clupeidae, Cyprinidae, Alestidae, Claroteidae, Clariidae, Malapteruridae, Mochokidae, Poeciliidae, Latidae, and Mastacembelidae) collected. Conclusion. Our records substantially expand the known range of fish species in a range of habitats. As numerous specimens are hard to assign to nominal species, a taxonomic revision of a number of genera is underway. It should take into account intraspecific geographic variation.