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The origin of introduced Nile tilapia stocks in sub-Saharan Africa is largely unknown. In this study, the potential of monogeneans as a biological tag and magnifying glass is tested to reveal their hosts' stocking history. The monogenean gill community of different Nile tilapia populations in sub-Saharan Africa was explored, and a phylogeographic analysis was performed based on the mitogenomes of four dactylogyrid species (Cichlidogyrus hani, C. sclerosus, C. thurstonae, and Scutogyrus longicornis). Our results encourage the use of dactylogyrids as biological tags. The magnifying glass hypothesis is only confirmed for C. thurstonae, highlighting the importance of the absence of other potential hosts as prerequisites for a parasite to act as a magnifying glass. With the data generated here, we are the first to extract mitogenomes from individual monogeneans and to perform an upscaled survey of the comparative phylogeography of several monogenean species with unprecedented diagnostic resolution. The Special Research Fund of Hasselt University finances MG (7NI02), AJCL (BOF19OWB02) and MPMV (BOF20TT06). Molecular work was funded through research grant 1513419 N of the Research Foundation—Flanders (FWO-Vlaanderen), and part of leading to results presented in this publication was carried out with infrastructure funded by the European Marine Biological Research Centre (EMBRC) Belgium, FWO-Vlaanderen project GOH3817N. The field expedition to the DRC was partly financed by the Stichting ter Bevordering van het Biodiversiteitsonderzoek in Afrika (SBBOA) fund.the research We appreciate the help of the people involved in the field work and sampling procedure. Furthermore, we would like to thank Christopher Laumer (EMBL-European Bioinformatics Institute, UK) and Gontran Sonet (Royal Belgium Institute of Natural Sciences, Belgium) for providing us with the extraction protocol and R scripts, respectively. We could also like to acknowledge Wim Bert, Christoph Hahn, Nikol Kmentova, ´ Jos Snoeks, and Carl Vangestel for suggestions on an earlier version of this manuscript. Finally, we would like to thank Ria Vanderspikken (Hasselt University, Belgium) and Natascha Steffanie (Hasselt University, Belgium) for their technical support in the laboratory

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.

The genus Clinostomoides Dollfus, 1950 was erected to accommodate a single worm from Ardea goliath sampled in the Belgian Congo. The specimen was distinguished from other clinostomids by its large size and posterior genitalia. In the following years, metacercariae of Clinostomoides brieni, have been described in Clarias spp. in southern and western Africa. A few authors have referred to Clinostomum brieni, but all such usages appear to be lapsus calami, and the validity of Clinostomoides remains widely accepted. In this study our aim was: position C. brieni among the growing clinostomids molecular database, and redescribe the species with emphasis on characters that have emerged as important in recent work. We sequenced two nuclear (partial 18S and ITS) and one mitochondrial marker (partial cytochrome c oxidase I) and studied morphology in metacercariae from hosts and localities likely to harbour the type species (Clarias spp., Democratic Republic of the Congo, South Africa). Phylogenetic analysis shows C. brieni belongs within Clinostomum Leidy, 1856. We therefore transfer C. brieni to Clinostomum, amend the diagnosis for the genus Clinostomum and provide a critical analysis of other species in Clinostomoides, all of which we consider species inquirendae, as they rest on comparisons of different developmental stages.

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.