Explosive networking: The role of adaptive host radiations and ecological opportunity in a species-rich host-parasite assembly.

Many species-rich ecological communities emerge from adaptive radiation events. Yet the effects of adaptive radiation on community assembly remain poorly understood. Here, we explore the well-documented radiations of African cichlid fishes and their interactions with the flatworm gill parasites Cichlidogyrus spp., including 10,529 reported infections and 477 different host-parasite combinations collected through a survey of peer-reviewed literature. We assess how evolutionary, ecological, and morphological parameters determine host-parasite meta-communities affected by adaptive radiation events through network metrics, host repertoire measures, and network link prediction. The hosts' evolutionary history mostly determined host repertoires of the parasites. Ecological and evolutionary parameters predicted host-parasite interactions. Generally, ecological opportunity and fitting have shaped cichlid-Cichlidogyrus meta-communities suggesting an invasive potential for hosts used in aquaculture. Meta-communities affected by adaptive radiations are increasingly specialised with higher environmental stability. These trends should be verified across other systems to infer generalities in the evolution of species-rich host-parasite networks.

Molecular Effects of Silver Nanoparticles on Monogenean Parasites: Lessons from Caenorhabditis elegans.

The mechanisms of action of silver nanoparticles (AgNPs) in monogenean parasites of the genus Cichlidogyrus were investigated through a microarray hybridization approach using genomic information from the nematode Caenorhabditis elegans. The effects of two concentrations of AgNPs were explored, low (6 µg/L Ag) and high (36 µg/L Ag). Microarray analysis revealed that both concentrations of AgNPs activated similar biological processes, although by different mechanisms. Expression profiles included genes involved in detoxification, neurotoxicity, modulation of cell signaling, reproduction, embryonic development, and tegument organization as the main biological processes dysregulated by AgNPs. Two important processes (DNA damage and cell death) were mostly activated in parasites exposed to the lower concentration of AgNPs. To our knowledge, this is the first study providing information on the sub-cellular and molecular effects of exposure to AgNPs in metazoan parasites of fish.

Responses of monogenean species to variations in abiotic parameters in tilapiculture.

Fish farming is becoming an increasingly popular agricultural activity, and water quality in these environments is a major concern. Fish parasites, such as monogeneans, respond to changes in abiotic conditions, either with an increase or decrease in population. This study aimed to identify gill monogeneans and analyse their relationships with abiotic factors during the ontogenetic development of Nile tilapia over the fish culture cycle in Mato Grosso do Sul, Brazil. Fish were sampled monthly for eight months, and a total of 200 fish were collected. The physical and chemical water parameters were measured and correlated with the abundance of each monogenean species. Over the fish culture cycle, the physical and chemical parameters fluctuated, and the water quality decreased. The parasites found included Cichlidogyrus tilapiae, Cichlidogyrus thurstonae, Cichlidogyrus sclerosus, Cichlidogyrus halli and Scutogyrus longicornis. The abundances of all species showed significant differences during ontogenetic development (body size) and C. tilapiae, C. sclerosus, C. thurstonae and S. longicornis were correlated with changes in abiotic conditions. However, C. halli was not significantly correlated with any of the evaluated physical or chemical parameters. Understanding how different monogenean species respond to changes in the physical and chemical parameters of water during a production cycle can prevent peaks in abundance and subsequent sanitary problems.

The first next-generation sequencing approach to the mitochondrial phylogeny of African monogenean parasites (Platyhelminthes: Gyrodactylidae and Dactylogyridae).

BACKGROUND: Monogenean flatworms are the main ectoparasites of fishes. Representatives of the species-rich families Gyrodactylidae and Dactylogyridae, especially those infecting cichlid fishes and clariid catfishes, are important parasites in African aquaculture, even more so due to the massive anthropogenic translocation of their hosts worldwide. Several questions on their evolution, such as the phylogenetic position of Macrogyrodactylus and the highly speciose Gyrodactylus, remain unresolved with available molecular markers. Also, diagnostics and population-level research would benefit from the development of higher-resolution genetic markers. We aim to offer genetic resources for work on African monogeneans by providing mitogenomic data of four species (two belonging to Gyrodactylidae, two to Dactylogyridae), and analysing their gene sequences and gene order from a phylogenetic perspective. RESULTS: Using Illumina technology, the first four mitochondrial genomes of African monogeneans were assembled and annotated for the cichlid parasites Gyrodactylus nyanzae, Cichlidogyrus halli, Cichlidogyrus mbirizei (near-complete mitogenome) and the catfish parasite Macrogyrodactylus karibae (near-complete mitogenome). Complete nuclear ribosomal operons were also retrieved, as molecular vouchers. The start codon TTG is new for Gyrodactylus and for Dactylogyridae, as is the incomplete stop codon TA for Dactylogyridae. Especially the nad2 gene is promising for primer development. Gene order was identical for protein-coding genes and differed between the African representatives of these families only in a tRNA gene transposition. A mitochondrial phylogeny based on an alignment of nearly 12,500 bp including 12 protein-coding and two ribosomal RNA genes confirms that the Neotropical oviparous Aglaiogyrodactylus forficulatus takes a sister group position with respect to the other gyrodactylids, instead of the supposedly 'primitive' African Macrogyrodactylus. Inclusion of the African Gyrodactylus nyanzae confirms the paraphyly of Gyrodactylus. The position of the African dactylogyrid Cichlidogyrus is unresolved, although gene order suggests it is closely related to marine ancyrocephalines. CONCLUSIONS: The amount of mitogenomic data available for gyrodactylids and dactylogyrids is increased by roughly one-third. Our study underscores the potential of mitochondrial genes and gene order in flatworm phylogenetics, and of next-generation sequencing for marker development for these non-model helminths for which few primers are available.

Expression of immune-related genes of Nile tilapia Oreochromis niloticus after Gyrodactylus cichlidarum and Cichlidogyrus sclerosus infections demonstrating immunosupression in coinfection.

Gyrodactylus cichlidarum and Cichlidogyrus sclerosus, two monogenean ectoparasite species commonly found on the body surface and gills of Nile tilapia (Oreochromis niloticus) respectively, inflicted considerable economic losses in intensive tilapia farming. In order to explore the immune response of tilapia against these two species of monogeneans, expression patterns of five immune-related genes were studied after singular G. cichlidarum or C. sclerosus infection and their coinfection. The transcription levels of IL-1ß were up-regulated in the skin after G. cichlidarum infection, reaching a peak at day 5 PI, and in the gills after C. sclerosus infection (peaking at day 8 PI), with significant elevation only detected in the gills after high-dose C. sclerosus infection. A trend favoring increased gill TNF-α expression at day 8 PI of C. sclerosus infection was statistically significant only in the low-dose infection group. TNF-α expression in the skin did not change significantly after G. cichlidarum infection. TGF-ß had extremely up-regulated expressions in the gills at day 8 PI after both high- and low-dose C. sclerosus infections, but its significantly promoted expression in the skin was observed only after infection of high-dose G. cichlidarum. Significantly increased expressions of HSP70 and COX-2 in the skin were detected after high-dose G. cichlidarum infections. In comparison to singular infection with either G. cichlidarum or C. sclerosus, concurrent infection resulted in significantly advanced expression of TGF-ß in both skin and gills, and lower expressions at day 8 PI, and similar patterns were observed in the expression of IL-1ß and TNF-α in the gills. G. cichlidarum infection on the body surface significantly down-regulated the expressions of TNF-α, TGF-ß and COX-2 in the gills. In addition, the intensity of G. cichlidarum was significantly positively correlated with that of C. sclerosus (correlation index 0.922, p = 0.000) at day 2 PI under concurrent infection. These results contribute to the understanding of mucosal immunity of fish against monogenean infection, particularly when two monogenean species infect concurrently.

Influence of preservative and mounting media on the size and shape of monogenean sclerites.

Based on Cichlidogyrus sp. (Monogenea, Ancyrocephalidae) specimens from Hemichromis sp. hosts, we tested the influence of different methods to fix/preserve samples/specimens [frozen material, alcohol or formalin preserved, museum process for fish preservation (fixed in formalin and preserved in alcohol)] and different media used to mount the slides [tap water, glycerin ammonium picrate (GAP), Hoyer's one (HM)] on the size/shape of sclerotized parts of monogenean specimens. The results show that the use of HM significantly increases the size of haptoral sclerites [marginal hooks I, II, IV, V, and VI; dorsal bar length, width, distance between auricles and auricle length, ventral bar length and width], and changes their shape [angle opening between shaft and guard (outer and inner roots) in both ventral and dorsal anchors, ventral bar much wider, dorsal one less curved]. This influence seems to be reduced when specimens/samples are fixed in formalin. The systematics of Monogenea being based on the size and shape of their sclerotized parts, to prevent misidentifications or description of invalid new species, we recommend the use of GAP as mounting medium; Hoyer's one should be restricted to monogenean specimens fixed for a long time which are more shrunken.

Gill monogeneans of Oreochromis niloticus (Linnaeus, 1758) and Oreochromis leucostictus (Trewavas, 1933) in Lake Naivasha, Kenya.

An investigation of gill monogeneans from the Nile tilapia Oreochromis niloticus and the blue spotted tilapia Oreochromis leucostictus (50 individuals per species) was done between the months of November 2014 to February 2015 in Lake Naivasha, Kenya. Standard parasitological procedures were used to examine fish gills for the presence of monogeneans. The observed monogeneans were collected, preliminarily identified using identification keys, quantified and fixed in 4 % formalin for morphological studies and absolute ethanol for molecular studies. Four parasite species comprising of three species of the genus Cichlidogyrus and one species of the genus Scutogyrus were recovered. Cichlidogyrus sclerosus and Cichlidogyrus tilapiae infested both fish species but the C. sclerosus was most prevalent in O. leucostictus (Prevalence (P) = 100 %, Mean intensity (MI) = 3.4) and C. tilapiae in O. niloticus (P = 8 %, MI = 4). Cichlidogyrus tilapiae had a P = 12 % and MI = 5.0 and a P = 6 % and MI = 3.0 in O. niloticus and O. leucostictus, respectively. Cichlidogyrus halli (P = 4 %, MI = 15.5) and Scutogyrus gravivaginus (P = 2 %, MI = 1.0) were only found in O. leucostictus. This is the first time that these monogeneans have been identified from Lake Naivasha, Kenya, presenting new geographical records. It was concluded that Ancyrocephalids (Cichilidogyrus spp.) dominated the two cichlid fish species in Lake Naivasha, Kenya.

Parasites (Monogenea) of tilapias Oreochromis niloticus and Coptodon rendalli (Cichlidae) in a river spring in Brazil.

In the present study, we examined 30 individuals of introduced African cichlids, Oreochromis niloticus and Coptodon rendalli, collected in a river spring of the Pardo River, Paranapanema River basin, southeastern Brazil. Based on morphological and molecular analyses of the partial LSU rDNA gene, we identified four species of monogeneans, Cichlidogyrus tilapiae, C. thurstonae, C. mbirizei, and Scutogyrus longicornis on the gills of O. niloticus, whereas individuals of C. rendalli were infested only with C. papernastrema. This is the first record of C. mbirizei and C. papernastrema in tilapias from Brazil. The ecological consequences of the introduction of exotic species of tilapia such as O. niloticus and C. rendalli along with their monogenean parasites in a wild environment represented by a river spring are discussed. Our new molecular data on Cichlidogyrus and Scutogyrus contribute to the investigation of the phylogenetic interrelationships of these widely distributed genera of monogeneans since their species composition is still unsettled.

Title: Parasites (Monogenea) des tilapias Oreochromis niloticus et Coptodon rendalli (Cichlidae) dans une source au Brésil. Abstract: Dans la présente étude, nous avons examiné 30 individus de cichlidés africains introduits, Oreochromis niloticus et Coptodon rendalli, collectés dans une source fluviale du fleuve Pardo, bassin du fleuve Paranapanema, dans le sud-est du Brésil. Sur la base d'analyses morphologiques et moléculaires du gène partiel de l'ADNr LSU, nous avons identifié quatre espèces de monogènes, Cichlidogyrus tilapiae, C. thurstonae, C. mbirizei et Scutogyrus longicornis sur les branchies d'O. niloticus, alors que les individus de C. rendalli étaient infestés uniquement par C. papernastrema. Il s'agit du premier signalement de C. mbirizei et C. papernastrema chez des tilapias du Brésil. Les conséquences écologiques de l'introduction d'espèces exotiques de tilapia telles que O. niloticus et C. rendalli ainsi que leurs monogènes parasites dans un environnement sauvage représenté par une source fluviale sont discutées. Nos nouvelles données moléculaires sur Cichlidogyrus et Scutogyrus contribuent à l'étude des interrelations phylogénétiques de ces genres de monogènes largement distribués puisque leur composition spécifique est encore incertaine.

High prevalence and mean intensity of trichodinids and monogeneans on Nile tilapia (Oreochromis niloticus) in Indonesian hatcheries.

This study describes recent infestations of ectoparasites on tilapia fingerlings in several hatcheries. High mortality of fingerlings and juvenile fish often occurs in hatcheries but the causative agents remain unclear. Nile tilapia fingerlings were obtained from three hatcheries in South Sulawesi, Indonesia. The fish were euthanized in clove oil, length and weight were measured, and the whole body and gills were examined for ectoparasites. Trichodinids were silver stained with 2% AgNO3, whereas monogeneans were placed in glycerin-alcohol and examined under a microscope. Other protozoans were placed directly on fresh mounts and identified under a compound microscope. The monogenean parasites were cleaned and placed in a microtube containing 70% alcohol for molecular identification. Trichodinid (Trichodina centrostrigeata, T. magna, Paratrichodina africana, Trichodinella sp., and Tripartiella sp.), and monogenean (Gyrodactylus sp. and Cichlidogyrus spp.) parasites were found in highly prevalent. The Cichlidogyrus specimens were morphologically identified as C. sclerosus, C. halli, and C. thurston, but only C. sclerosus and C. halli were confirmed based on partial 28S rRNA. The Gyrodactylus specimens were morphologically identified as G. cichlidarum and confirmed based on the ITS region. This is the first time that T. centrostrigeata and P. africana have been confirmed on fish from hatcheries and is also the first time that C. sclerosus, C. halli, and G. cichlidarum have been verified in Indonesian hatcheries based on a molecular technique. The high infestation rates of these parasites were likely caused by rearing the fish under stressful conditions, indicating low biosecurity and poor health management practices in the aquaculture facilities.

First record of monogenean fish parasites in the Upper Lufira River Basin (Democratic Republic of Congo): dactylogyrids and gyrodactylids infesting Oreochromis mweruensis, Coptodon rendalli and Serranochromis macrocephalus (Teleostei: Cichlidae).

BACKGROUND: Monogenean parasites have never been formally reported on fishes from the Lufira River Basin. In this context, we decided to record the monogenean parasite fauna of three cichlid species found in the Upper Lufira River Basin for the first time by inventorizing their diversity (species composition) and analysing their infection parameters (prevalence, mean intensity and abundance). METHODS: The African cichlid fishes Oreochromis mweruensis, Coptodon rendalli and Serranochromis macrocephalus were selected for the study, given their economic value and their abundance in the Upper Lufira River Basin. Monogeneans were isolated from the gills and stomach, mounted on glass slides with either Hoyer's medium or ammonium picrate-glycerin for identification under a stereomicroscope, based on morphological analysis of genital and haptoral hard parts. Indices of diversity and infections parameters were calculated. RESULTS: A total of 13 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 identified. A species richness (S) of 10 for O. mweruensis, S = 6 for C. rendalli and S = 2 for S. macrocephalus was recorded. Five parasite species were reported to be common amongst O. mweruensis and C. rendalli. According to cichlid species, the most prevalent parasite species was C. halli (prevalence [P] = 80.9%) on O. mweruensis, C. dossoui (P = 92.9%) on C. rendalli and C. karibae and C. zambezensis (both P = 9.1%) on S. macrocephalus. The parasite species with the highest mean intensity (MI) were G. nyanzae (MI = 8.7) on O. mweruensis, C. papernastrema (MI = 17.1) on C. rendalli and C. karibae (MI = 15) on S. macrocephalus. The findings indicate 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 all of them as they are recorded for the first time in the Lufira River Basin. CONCLUSIONS: This study highlighted the richness of monogenean communities in the Upper Lufira River Basin and is a starting point for future helminthological studies, such as on the use of fish parasites as indicators of anthropogenic impacts.

Sharing is caring? Barcoding suggests co-introduction of dactylogyrid monogeneans with Nile tilapia and transfer towards native tilapias in sub-Saharan Africa.

Invasive Nile tilapias negatively impact native tilapia species through hybridisation and competition. However, the co-introduction of parasites with Nile tilapia, and subsequent changes in parasite communities, are scarcely documented. Monogeneans are known pathogens of cultured Nile tilapia, although little is known about their fate once Nile tilapias establish in new ecosystems. We investigate the parasitological consequences of Nile tilapia introduction on native tilapias in basins in Cameroon, the Democratic Republic of the Congo (DRC), and Zimbabwe, focusing on ectoparasitic dactylogyrids (Monogenea). Using the mitochondrial cytochrome oxidase c subunit I (COI) and nuclear 18S-internal transcribed spacer 1 (18S-ITS1) rDNA region of 128 and 166 worms, respectively, we evaluated transmission of several dactylogyrid species. Parasite spillover from Nile tilapia was detected for Cichlidogyrus tilapiae to Coptodon guineensis in Cameroon, Cichlidogyrus thurstonae to Oreochromis macrochir in the DRC, and Cichlidogyrus halli and C. tilapiae to Coptodon rendalli in Zimbabwe. Parasite spillback to Nile tilapia was detected for Cichlidogyrus papernastrema and Scutogyrus gravivaginus from Tilapia sparrmanii and Cichlidogyrus dossoui from C. rendalli or T. sparrmanii in the DRC, and Cichlidogyrus chloeae from Oreochromis cf. mortimeri and S. gravivaginus from O. macrochir in Zimbabwe. 'Hidden' transmissions (i.e. transmission of certain parasite lineages of species that are naturally present on both alien and native hosts) were detected for C. tilapiae and Scutogyrus longicornis between Nile tilapia and Oreochromis aureus and C. tilapiae between Nile tilapia and Oreochromis mweruensis in the DRC, and Cichlidogyrus sclerosus and C. tilapiae between Nile tilapia and O. cf. mortimeri in Zimbabwe. A high density of Nile tilapia occurring together with native tilapias, and the broad host range and/or environmental tolerance of the transmitted parasites, are proposed as factors behind parasite transmission through ecological fitting. However, continuous monitoring and the inclusion of environmental variables are necessary to understand the long-term consequences of these transmissions on native tilapias and to elucidate other underlying factors influencing these transmissions.

Contrasting Host-Parasite Population Structure: Morphology and Mitogenomics of a Parasitic Flatworm on Pelagic Deepwater Cichlid Fishes from Lake Tanganyika.

Little phylogeographic structure is presumed for highly mobile species in pelagic zones. Lake Tanganyika is a unique ecosystem with a speciose and largely endemic fauna famous for its remarkable evolutionary history. In bathybatine cichlid fishes, the pattern of lake-wide population differentiation differs among species. We assessed the congruence between the phylogeographic structure of bathybatine cichlids and their parasitic flatworm Cichlidogyrus casuarinus to test the magnifying glass hypothesis. Additionally, we evaluated the use of a PoolSeq approach to study intraspecific variation in dactylogyrid monogeneans. The lake-wide population structure of C. casuarinus ex Hemibates stenosoma was assessed based on a portion of the cox1 gene combined with morphological characterisation. Additionally, intraspecific mitogenomic variation among 80 parasite samples from one spatially constrained metapopulation was assessed using shotgun NGS. While no clear geographic genetic structure was detected in parasites, both geographic and host-related phenotypic variation was apparent. The incongruence with the genetic north-south gradient observed in H. stenosoma may be explained by the broad host range of this flatworm including eupelagic bathybatine host species that form panmictic populations across the lake. In addition, we present the first parasite mitogenome from Lake Tanganyika and propose a methodological framework for studying the intraspecific mitogenomic variation of dactylogyrid monogeneans.

Six new species of Cichlidogyrus Paperna, 1960 (Platyhelminthes: Monogenea) from the gills of cichlids (Teleostei: Cichliformes) from the Lomami River Basin (DRC: Middle Congo).

BACKGROUND: Monogenea van Beneden, 1858 is a group of parasitic flatworms, commonly found infecting bony fish. Several genera, such as Cichlidogyrus Paperna, 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 of Cichlidogyrus in the south of the Democratic Republic of the Congo (DRC) on three cichlid species: Orthochromis sp. 'Lomami', Serranochromis cf. macrocephalus, and Tilapia sparrmanii Smith, 1840. METHODS: Specimens of Cichlidogyrus were 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. RESULTS: In total, six species of Cichlidogyrus were found, all new to science: C. bulbophallus n. sp. and C. pseudozambezensis n. sp. on S. cf. macrocephalus, C. flagellum n. sp. and C. lobus n. sp. on T. sparrmanii, C. ranula n. sp. on S. cf. macrocephalus and Orthochromis sp. 'Lomami', and C. maeander n. sp. found on Orthochromis sp. 'Lomami' and T. sparrmanii. The first four species are considered to be strict specialists, C. ranula n. sp. an intermediate generalist and C. maeander n. 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. CONCLUSIONS: Serranochromis cf. macrocephalus and Orthochromis sp. 'Lomami' can harbour respectively three and two species of Cichlidogyrus, all described in this study. Tilapia sparrmanii can 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.

Underexplored diversity of gill monogeneans in cichlids from Lake Tanganyika: eight new species of Cichlidogyrus Paperna, 1960 (Monogenea: Dactylogyridae) from the northern basin of the lake, with remarks on the vagina and the heel of the male copulatory organ.

BACKGROUND: Lake Tanganyika harbours the most diverse cichlid assemblage of the Great African Lakes. Considering its cichlid flocks consist of approximately 250 endemic species, we can hypothesize a high species-richness in their often quite host-specific monogenean ectoparasites belonging to Cichlidogyrus Paperna, 1960. Yet, only 24 species were described from Tanganyikan hosts and some host tribes have never been investigated for monogeneans. This study presents the first parasitological examination of species of the tribes Cyprichromini (Cyprichromis microlepidotus (Poll, 1956)), Eretmodini (Eretmodus marksmithi Burgess, 2012 and Tanganicodus irsacae Poll, 1950) and Ectodini (Aulonocranus dewindti (Boulenger, 1899)). Specimens of the ectodine Ophthalmotilapia nasuta (Poll & Matthes, 1962) from which four Cichlidogyrus spp. have been previously described from more southern localities were also studied. Further, we discuss the haptor configuration in Tanganyikan Cichlidogyrus spp. and highlight the morphological diversity of the vagina, and that of the heel, a sclerotized part of the male copulatory organ, absent in some species of Cichlidogyrus. METHODS: Cichlidogyrus spp. were isolated from gills and fixed using GAP. Haptoral and genital hard parts were measured and drawn by means of a phase contrast microscopic examination. RESULTS: We describe eight new species: Cichlidogyrus milangelnari n. sp. on C. microlepidotus; C. jeanloujustinei n. sp. on E. marksmithi; C. evikae n. sp. on T. irsacae; C. aspiralis n. sp., C. glacicremoratus n. sp. and C. rectangulus n. sp. on O. nasuta; and C. pseudoaspiralis n. sp. and C. discophonum n. sp. on A. dewindti. Three haptoral morphotypes were recognized among the new species. Species of Cichlidogyrus from closely related hosts exhibited the same morphotypes. Geographical variation in Cichlidogyrus spp. fauna as observed in O. nasuta and three morphotypes were distinguished. Finally, we listed 111 Cichlidogyrus species, of which 27 and three Tanganyikan species lack sclerotized vagina and heel, respectively, just like 19 and seven species outside of the lake. CONCLUSIONS: Haptoral and genital features in the Tanganyikan Cichlidogyrus fauna reflect the phylogenetic relationships of their cichlid hosts. It seems that several lineages of Cichlidogyrus spp. exist in Lake Tanganyika but further studies are necessary to confirm this hypothesis and answer questions related to Lake Tanganyika and its cichlids.

First insights into the diversity of gill monogeneans of 'Gnathochromis' and Limnochromis (Teleostei, Cichlidae) in Burundi: do the parasites mirror host ecology and phylogenetic history?

Monogenea is one of the most species-rich groups of parasitic flatworms worldwide, with many species described only recently, which is particularly true for African monogeneans. For example, Cichlidogyrus, a genus mostly occurring on African cichlids, comprises more than 100 nominal species. Twenty-two of these have been described from Lake Tanganyika, a famous biodiversity hotspot in which many vertebrate and invertebrate taxa, including monogeneans, underwent unique and spectacular radiations. Given their often high degrees of host specificity, parasitic monogeneans were also used as a potential tool to uncover host species relationships. This study presents the first investigation of the monogenean fauna occurring on the gills of endemic 'Gnathochromis' species along the Burundese coastline of Lake Tanganyika. We test whether their monogenean fauna reflects the different phylogenetic position and ecological niche of 'Gnathochromis' pfefferi and Gnathochromis permaxillaris. Worms collected from specimens of Limnochromis auritus, a cichlid belonging to the same cichlid tribe as G. permaxillaris, were used for comparison. Morphological as well as genetic characterisation was used for parasite identification. In total, all 73 Cichlidogyrus individuals collected from 'G.' pfefferi were identified as C. irenae. This is the only representative of Cichlidogyrus previously described from 'G.' pfefferi, its type host. Gnathochromis permaxillaris is infected by a species of Cichlidogyrus morphologically very similar to C. gillardinae. The monogenean species collected from L. auritus is considered as new for science, but sample size was insufficient for a formal description. Our results confirm previous suggestions that 'G.' pfefferi as a good disperser is infected by a single monogenean species across the entire Lake Tanganyika. Although G. permaxillaris and L. auritus are placed in the same tribe, Cichlidogyrus sp. occurring on G. permaxillaris is morphologically more similar to C. irenae from 'G.' pfefferi, than to the Cichlidogyrus species found on L. auritus. Various evolutionary processes, such as host-switching or duplication events, might underlie the pattern observed in this particular parasite-host system. Additional samples for the Cichlidogyrus species occuring on G. permaxillaris and L. auritus are needed to unravel their evolutionary history by means of (co-)phylogenetic analyses.

Gill monogeneans of Nile tilapia (Oreochromis niloticus) and red hybrid tilapia (Oreochromis spp.) from the wild and fish farms in Perak, Malaysia: infection dynamics and spatial distribution.

Tilapia is one of the commercially important fish in Malaysia as well as in other parts of the world. An understanding of monogenean infection dynamics in tilapia fish may assist us in searching for some intervention measures in reducing the loss of fish caused by parasitic diseases. The present study aimed (1) to compare infection level of monogeneans between the wild and cultured Oreochromis niloticus, and between the cultured O. niloticus and cultured red hybrid tilapia, and (2) to examine the spatial distribution of monogenean species over the gills of the different host species. From a total of 75 fish specimens, six species of monogeneans from two genera: Cichlidogyrus (C. halli, C. mbirizei, C. sclerosus, C. thurstonae, C. tilapiae) and Scutogyrus (S. longicornis) were identified. Data showed that the infection level of cultured O. niloticus was higher than that of the wild O. niloticus, however, the former was lower than that of the cultured red hybrid tilapia. Higher species richness of monogeneans was observed in the cultured red hybrid tilapia as compared to the others. Results for spatial distribution showed that the monogeneans have no preference on the left or right sides of the gills. However, C. halli, C. mbirizei, and C. tilapiae showed preferences on specific gill arches in the cultured O. niloticus and red hybrid tilapia. In general, the gill arch IV harboured the least number of monogeneans. The susceptibility of monogenean infection between the different types of tilapia is discussed.

Deep-water parasite diversity in Lake Tanganyika: description of two new monogenean species from benthopelagic cichlid fishes.

BACKGROUND: Lake Tanganyika is the world's second deepest lake. Its diverse cichlid assemblage offers a unique opportunity for studying a deep-water host-parasite model in freshwater. Low host specificity and a broad host range including representatives of the Bathybatini tribe in the only monogenean parasite described from this habitat, Cichlidogyrus casuarinus Pariselle, Muterezi Bukinga & Vanhove, 2015 suggest a link between lower specificity and lower host density. Conversely, high host specificity and species richness are reported for monogeneans of the lake's littoral cichlids. We further investigated whether the deep-water environment in Lake Tanganyika is really monogenean species-depauperate by investigating the monogenean fauna of Trematocara unimaculatum (a representative of the tribe Trematocarini, the sister lineage of the Bathybatini) and Benthochromis horii, a member of the tribe Benthochromini, found in the same deep-water habitat as the already known hosts of C. casuarinus. METHODS: Sclerotised structures of the collected monogenean individuals were characterised morphologically using light microscopy and morphometrics. RESULTS: Both examined cichlid species are infected by a single monogenean species each, which are new to science. They are described as Cichlidogyrus brunnensis n. sp., infecting T. unimaculatum, and Cichlidogyrus attenboroughi n. sp., parasitising on B. horii. Diagnostic characteristics include the distal bifurcation of the accessory piece in C. brunnensis n. sp. and the combination of long auricles and no heel in C. attenboroughi n. sp. In addition C. brunnensis n. sp. does not resemble C. casuarinus, the only species of Cichlidogyrus thus far reported from the Bathybatini. Also Cichlidogyrus attenboroughi n. sp. does not resemble any of the monogenean species documented from the pelagic zone of the lake and is among the few described species of Cichlidogyrus without heel. CONCLUSIONS: As two new and non-resembling Cichlidogyrus species are described from T. unimaculatum and B. horii, colonisation of the deep-water habitat by more than one morphotype of Cichlidogyrus is evident. Based on morphological comparisons with previously described monogenean species, parasite transfers with the littoral zone are possible. Therefore, parasites of pelagic cichlids in the lake do not seem to only mirror host phylogeny and the evolutionary history of this host-parasite system merits further attention.

First report of monogenean flatworms from Lake Tana, Ethiopia: gill parasites of the commercially important Clarias gariepinus (Teleostei: Clariidae) and Oreochromis niloticus tana (Teleostei: Cichlidae).

BACKGROUND: Lake Tana is the largest lake in Ethiopia and the source of the Blue Nile. The lake harbours unique endemic cyprinid fish species, as well as the commercially important endemic Nile tilapia subspecies Oreochromis niloticus tana and the North African catfish Clarias gariepinus. Its endemicity, especially within the Labeobarbus radiation, its conservation importance and its economic indispensability attract scientific interest to the lake's ichthyofauna. Fish parasites of Lake Tana, however, are hitherto poorly known, and no formal report exists on its monogenean flatworms. For sustainable aquaculture and fisheries development, it is essential to study monogenean fish parasites in these economically most important fish species. Moreover, it remains to be verified whether this unique ecosystem and its endemicity gave rise to a distinct parasite fauna as well. RESULTS: Nile tilapia and North African catfish hosts were collected from Lake Tana in 2013. Nine species of monogenean parasites of two orders, Gyrodactylidea Bychowsky, 1937 and Dactylogyridea Bychowsky, 1937, were recovered. Gyrodactylus gelnari Prikrylová, Blazek & Vanhove, 2012, Macrogyrodactylus clarii Gussev, 1961, Quadriacanthus aegypticus El-Naggar & Serag, 1986 and two undescribed Quadriacanthus species were recovered from C. gariepinus. Oreochromis niloticus tana hosted Cichlidogyrus cirratus Paperna, 1964, C. halli (Price & Kirk, 1967), C. thurstonae Ergens, 1981 and Scutogyrus longicornis (Paperna & Thurston, 1969). CONCLUSIONS: Except for M. clarii, all species represent new records for Ethiopia. This first study on the monogenean fauna of Lake Tana revealed that the lake's North African catfish, as well as its endemic Nile tilapia subspecies, harbour parasites that are known from these host species elsewhere in Africa.