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.

Somewhere I belong: phylogeny and morphological evolution in a species-rich lineage of ectoparasitic flatworms infecting cichlid fishes.

A substantial portion of biodiversity has evolved through adaptive radiation. However, the effects of explosive speciation on species interactions remain poorly understood. Metazoan parasites infecting radiating host lineages could improve our knowledge because of their intimate host relationships. Yet limited molecular, phenotypic and ecological data discourage multivariate analyses of evolutionary patterns and encourage the use of discrete characters. Here, we assemble new molecular, morphological and host range data widely inferred from a species-rich lineage of parasites (Cichlidogyrus, Platyhelminthes: Monogenea) infecting cichlid fishes to address data scarcity. We infer a multimarker (28S/18S rDNA, ITS1, COI mtDNA) phylogeny of 58 of 137 species and characterize major lineages through synapomorphies inferred from mapping morphological characters. We predict the phylogenetic position of species without DNA data through shared character states, a morphological phylogenetic analysis, and a classification analysis with support vector machines. Based on these predictions and a cluster analysis, we assess the systematic informativeness of continuous characters, search for continuous equivalents for discrete characters, and suggest new characters for morphological traits not analysed to date. We also model the attachment/reproductive organ and host range evolution using the data for 136 of 137 described species and multivariate phylogenetic comparative methods (PCMs). We show that discrete characters not only can mask phylogenetic signals, but also are key for characterizing species groups. Regarding the attachment organ morphology, a divergent evolutionary regime for at least one lineage was detected and a limited morphological variation indicates host and environmental parameters affecting its evolution. However, moderate success in predicting phylogenetic positions, and a low systematic informativeness and high multicollinearity of morphological characters call for a revaluation of characters included in species characterizations.

Temporally consistent species differences in parasite infection but no evidence for rapid parasite-mediated speciation in Lake Victoria cichlid fish.

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.

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.

Four new species of Dactylogyrus Diesing, 1850 (Monogenea: Dactylogyridae) parasitising the gills of northern Moroccan Luciobarbus Heckel (Cyprinidae): morphological and molecular characterisation.

Four new species of Dactylogyrus Diesing, 1850 are described from the gills of three species of Luciobarbus Heckel collected from various hydrographical basins in northern Morocco: Dactylogyrus scorpius n. sp. from Luciobarbus rifensis Doadrio, Casal-Lopez & Yahyaoui; D. benhoussai n. sp. from L. moulouyensis Pellegrin; and D. varius n. sp. and D. falsiphallus n. sp. from L. maghrebensis Doadrio, Perea & Yahyaoui. The descriptions of the new species are confirmed by molecular data (partial 18S rDNA, ITS1, and partial 28S rDNA sequences). All four species belong to the group of Dactylogyrus species, possessing a cross-shaped ventral bar and a male copulatory organ composed of a loosely coiled copulatory tube and an accessory piece with a capsule-like base and recurved distal portion. Given the high shape variability of the haptoral anchors reported among specimens of D. varius n. sp., three morphological forms within this species (D. varius f. vulgaris, D. varius f. magnus, and D. varius f. dromedarius) are recognised. However, specimens belonging to D. benhoussai n. sp. and D. varius f. vulgaris were morphologically very similar and were discriminated with certainty, only when using molecular data.

A case of complete loss of gill parasites in the invasive cichlid Oreochromis mossambicus.

This study investigates the recent evolution of a rich parasite community associated with one of the world's most invasive species, the cichlid fish Oreochromis mossambicus. Populations from the species' native range (Mozambique) are compared to a population from New Caledonia (Wester Pacific), an island where the species was introduced in 1954. The results support the complete local extinction of the gill parasite community in the course of the invasion process. Up to six gill parasite species per locality were documented in the O. mossambicus native range, and previous surveys consistently reported at least one parasite species introduced along African cichlid species established out of Africa. The absence of parasites in New Caledonia is therefore exceptional. This can be attributed to local factors, such as a strong initial population bottleneck, the likely absence of multiple host introductions, and the frequent occurrence of brackish watersheds that might enhance the probability for natural deparasitation.

A comparative study of Ligophorus uruguayense and L. saladensis (Monogenea: Ancyrocephalidae) from Mugil liza (Teleostei: Mugilidae) in southern Brazil.

Representatives of Ligophorus Euzet et Suriano, 1977 were found on the gills of Mugil liza Valenciennes caught in southern Brazil. They were identified as Ligophorus uruguayense Failla Siquier et Ostrowski de Núñez, 2009 and Ligophorus saladensis Marcotegui et Martorelli, 2009, even though specific identification proved to be difficult due to inconsistencies in some diagnostic features reported for these two species. Therefore, a combined morphological and molecular approach was used to critically review the validity of these species, by means of phase contrast and confocal fluorescence microscopical examination of sclerotised hard parts, and assessing the genetic divergence between L. saladensis, L. uruguayense and their congeners using rDNA sequences. The main morphological differences between the two species relate to the shape of the accessory piece of the penis and the median process of the ventral bar. The accessory piece in L. uruguayense is shorter than in L. saladensis, has a cylindrical, convex upper lobe and straight lower lobe (vs with the distal tip of the lower lobe turning away from the upper lobe in the latter species). The ventral bar has a V-shaped anterior median part in L. uruguayense (vs U-shaped in L. saladensis). The two species are suggested to be part of a species complex together with L. mediterraneus Sarabeev, Balbuena et Euzet, 2005. We recommend to generalise such comparative assessment of species of Ligophorus for a reliable picture of the diversity and diversification mechanisms within the genus, and to make full use of its potential as an additional marker for mullet taxonomy and systematics.

Understanding the Influence of Host Radiation on Symbiont Speciation through Parasites of Species Flocks.

(Adaptive) radiations have attracted evolutionary biologists for a long time as ideal model systems to study patterns and processes of often rapid speciation. However, whereas a wealth of (sometimes already genome-scale) data is available for host radiations, very few studies target the patterns of diversification in their symbionts, even though they would be excellent models to study symbiont speciation. Our review summarizes what little is known about general patterns of symbiont diversification in often iconic adaptive host radiations and to what extent these patterns are dependent on the evolutionary trajectories of their hosts. We identify research gaps that need to be addressed in the future and discuss the potential of approaches not yet typically used in these study systems, such as epidemiological disease modeling and new omics technologies, for significantly advancing our understanding of these complex eco-evolutionary relationships.

Contrasting parasite communities among allopatric colour morphs of the Lake Tanganyika cichlid Tropheus.

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.

Ligophorus species (Monogenea: Ancyrocephalidae) from Mugil cephalus (Teleostei: Mugilidae) off Morocco with the description of a new species and remarks about the use of Ligophorus spp. as biological markers of host populations.

Gill monogenean species of Ligophorus Euzet et Suriano, 1977 were studied from the teleost Mugil cephalus Linneaus (Mugilidae) from the Mediterranean and Atlantic coasts of Morocco. We report the presence of L. mediterraneus from both the Mediterranean and Atlantic coast and L. cephali and L. maroccanus sp. n. from the Atlantic coast only. The latter species, which is described herein as new, resembles L. guanduensis but differs from this species mainly in having a shorter penis compared to the accessory piece, a proportionally longer extremity of the accessory piece and a less developed heel. The utility of Ligophorus spp. as markers of cryptic species of the complex M. cephalus is discussed in the context of species diversity and geographical distribution of these monogeneans on this host around the world. Presence of different species of Ligophorus on M. cephalus sensu stricto from the Atlantic and Mediterranean coast of Morocco demonstrates the usefulness of these species as fine resolution markers of genetic populations of their host, which are known to inhabit those coasts.

Checklist of the ancyrocephalids (Monogenea) parasitizing Tilapia species in Cameroon, with the description of three new species.

A checklist of the 23 species of Ancyrocephalidae (Monogenea) parasitizing Tilapia species sampled from both sides of the Cameroon Volcanic Line is provided. As already reported elsewhere, Cichlidogyrus aegypticus in Cameroon shows two different shape of its vagina, and C. tilapiae, C. arthracanthus and C. tiberianus were found on a wide range of host species. Among the 23 studied species of Monogenea, three are considered new species and are described herein: Cichlidogyrus berminensis n. sp. from Tilapia bemini, characterized by a short penis with marked narrow heel, a simple and straight accessory piece ending in a large hook, and a medium sized pair of uncinuli I; Cichlidogyrus gillesi n. sp. from Tilapia guineensis, characterized by a large and trapezoid heel of the penis and a S-shaped and wrinkle walled vagina; and Scutogyrus vanhovei n. sp. from Tilapia mariae, characterized by the presence of a distinct swollen portion of the penis. From a parasite's point of view, the CVL has no influence on species dispersion/distribution.

Biogeographical implications of Zambezian Cichlidogyrus species (Platyhelminthes: Monogenea: Ancyrocephalidae) parasitizing Congolian cichlids.

Fishes normally restricted to inland waters are valuable model systems for historical biogeography, inter alia, because of their limited dispersal abilities and concordance with the distribution patterns of other freshwater taxa (Zogaris et al. 2009). The comparison of fish species assemblages has been the major biogeographical tool for delineating African aquatic ecoregions as the fossil record is often meagre and merely offers complementary information. This is, for example, the case for the Zambezian and Congolian ichthyofaunal provinces, which display substantial contemporary fish diversity (Stewart 2001). Between both regions lies the Bangweulu-Mweru ecoregion (sensu Scott 2005), known for its high percentage of endemicity. Although hydrographically belonging to the Congo Basin, the Bangweulu-Mweru ecoregion has a high affinity with the Zambezi province (Scott 2005), due to historical river connections (Tweddle 2010). Studies comparing the Zambezi and Congo ichthyofaunal provinces are rare and hampered by lack of data from the Congo Basin. The latter harbours more than 1250 fish species (Snoeks et al. 2011) while in the Zambezi, only 120 freshwater fishes are found (Tweddle 2010). Indeed, species richness declines in all major African teleost families from the Congo Basin southwards, riverine haplochromine cichlids forming a notable exception to this rule (Joyce et al. 2005). Although it was hypothesized by Tweddle (2010) that the origin of many Zambezian fish species is in the Congo Basin, the haplochromines Serranochromis Regan, Sargochromis Regan, Pharyngochromis Greenwood and Chetia Trewavas, together forming the serranochromines, have their centre of diversity in the rivers of the Zambezian ichthyofaunal province (Joyce et al. 2005). Therefore, the biogeographical history of Cichlidae across the Zambezi- Congo watershed is not only key to cichlid biogeography on an African scale, but also complementary to biogeography of all other teleosts in the region. Yet, colonisation and speciation patterns are difficult to unravel due to complex hydrological history (Katongo et al. 2007; Schwarzer et al. 2012).

First Study on Gyrodactylus (Monogenea: Gyrodactylidae) in Morocco, with Description of a New Species from Luciobarbus pallaryi and Luciobarbus ksibi (Actinopterygii: Cyprinidae).

To date, 41 species of Gyrodactylus have been described from Africa. However, none of these have been reported in Morocco. After identifying and examining 738 cyprinid host specimens, 26 specimens belonging to Gyrodactylus were found to parasitize the gills of nine species of Luciobarbus, Carasobarbus, and Pterocapoeta. The current study provides new information about the presence of a new parasitic species in Morocco, the first to be characterized on a species level in the Maghreb region. It describes in detail 12 specimens of Gyrodactylus isolated from the gills of Luciobarbus pallaryi (Pellegrin, 1919) and Luciobarbus ksibi (Boulenger, 1905). Based on morphoanatomical observations, the characterization of the specimens collected indicates a species of Gyrodactylus that is new to science, described here as Gyrodactylus nyingiae n. sp. The new species is different from previously described gyrodactylids infecting African cyprinid hosts because it has a longer hamulus total length, a longer hamulus root, a downward projecting toe of the marginal hook, and a trapezium-shaped ventral bar membrane with a slightly striated median portion and small rounded anterolateral processes. This study increases the total number of Gyrodactylus spp. found in African cyprinids to four.

Monogeneans from Catfishes in Lake Tanganyika. II: New Infection Site, New Record, and Additional Details on the Morphology of the Male Copulatory Organ of Gyrodactylus transvaalensis Prudhoe and Hussey, 1977.

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.

All quiet on the western front? The evolutionary history of monogeneans (Dactylogyridae: Cichlidogyrus, Onchobdella) infecting a West and Central African tribe of cichlid fishes (Chromidotilapiini).

Owing to the largely unexplored diversity of metazoan parasites, their speciation mechanisms and the circumstances under which such speciation occurs - in allopatry or sympatry - remain vastly understudied. Cichlids and their monogenean flatworm parasites have previously served as a study system for macroevolutionary processes, e.g., for the role of East African host radiations on parasite communities. Here, we investigate the diversity and evolution of the poorly explored monogeneans infecting a West and Central African lineage of cichlid fishes: Chromidotilapiini, which is the most species-rich tribe of cichlids in this region. We screened gills of 149 host specimens (27 species) from natural history collections and measured systematically informative characters of the sclerotised attachment and reproductive organs of the parasites. Ten monogenean species (Dactylogyridae: Cichlidogyrus and Onchobdella) were found, eight of which are newly described and one redescribed herein. The phylogenetic positions of chromidotilapiines-infecting species of Cichlidogyrus were inferred through a parsimony analysis of the morphological characters. Furthermore, we employed machine learning algorithms to detect morphological features associated with the main lineages of Cichlidogyrus. Although the results of these experimental algorithms remain inconclusive, the parsimony analysis indicates that West and Central African lineages of Cichlidogyrus and Onchobdella are monophyletic, unlike the paraphyletic host lineages. Several instances of host sharing suggest occurrences of intra-host speciation (sympatry) and host switching (allopatry). Some morphological variation was recorded that may also indicate the presence of species complexes. We conclude that collection material can provide important insights on parasite evolution despite the lack of well-preserved DNA material.

Title: À l'Ouest, rien de nouveau ? L'histoire évolutive des monogènes (Dactylogyridae : Cichlidogyrus, Onchobdella) infectant une tribu de poissons cichlidés (Chromidotilapiini) d'Afrique occidentale et centrale. Abstract: En raison de la nature largement inexplorée de la diversité des parasites métazoaires, leurs mécanismes de spéciation et les circonstances dans lesquelles cette spéciation se produit­allopatrie ou sympatrie­restent très peu étudiés. Les cichlidés et leurs parasites Plathelminthes monogènes ont déjà servi de modèle pour l'étude des processus macro-évolutifs, par exemple pour le rôle des radiations d'hôtes de l'Afrique de l'Est sur les communautés de parasites. Ici, nous étudions la diversité et l'évolution des monogènes peu étudiées qui infestent une lignée de poissons cichlidés d'Afrique occidentale et centrale : les Chromidotilapiini, qui est la tribu de cichlidés la plus riche en espèces dans cette région. Nous avons examiné les branchies de 149 spécimens hôtes (27 espèces) provenant de musées d'histoire naturelle et mesuré systématiquement les caractères informatifs des pièces sclérifiées du hapteur et des organes copulateurs des parasites. Dix espèces de monogènes (Dactylogyridae : Cichlidogyrus et Onchobdella) ont été trouvées ; huit sont nouvelles pour la science et une est redécrite. Les positions phylogénétiques des espèces de Cichlidogyrus infectant les chromidotilapiines ont été déduites par une analyse de parcimonie des caractères morphologiques. En outre, nous avons utilisé des algorithmes d'apprentissage automatique pour détecter les caractéristiques morphologiques associées aux principales lignées de Cichlidogyrus. Bien que les résultats de ces algorithmes expérimentaux restent peu concluants, l'analyse de parcimonie indique que les lignées de Cichlidogyrus et d'Onchobdella de l'Afrique de l'Ouest et Central sont monophylétiques, contrairement aux lignées d'hôtes qui sont paraphylétiques. Plusieurs cas de partage d'hôtes suggèrent des occurrences de spéciation synxénique (sympatrie) et de changement d'hôte (allopatrie). Certaines variations morphologiques ont été enregistrées et peuvent également indiquer la présence de complexes d'espèces. Nous concluons donc que le matériel de collection peut fournir des informations importantes sur l'évolution des parasites malgré le manque d'ADN exploitable.

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.

Ancyrocephalidae (Monogenea) of Lake Tanganyika: II: description of the first Cichlidogyrus spp. parasites from Tropheini fish hosts (Teleostei, Cichlidae).

Although Lake Tanganyika hosts the most diverse endemic cichlid fish assemblage, its monogenean parasite fauna has hardly been documented. The cichlid tribe Tropheini has generated great interest because of its systematic position within the Haplochromini s.l. and its diversity in trophic morphology, reproductive behaviour and population structure. It has the potential to host a diverse Monogenea fauna. Here, we describe the first Cichlidogyrus spp.: Cichlidogyrus steenbergei sp. n., Cichlidogyrus irenae sp. n. and Cichlidogyrus gistelincki sp. n. The three host species, Limnotilapia dardennii, Ctenochromis horei and Gnathochromis pfefferi, are all infected by a single unique Cichlidogyrus sp. The genital and haptoral structure of the new species suggests a close relationship, which might mirror the close affinities between the hosts within the Tropheini. Based on haptoral configuration, the new species belong to a morphological group within the genus containing parasites both of West African cichlids and of Haplochromini, and hence, do not represent a new organisation of the attachment organ (as has recently been described of congeners infecting the ectodine cichlid Ophthalmotilapia).

Ancyrocephalidae (Monogenea) of Lake Tanganyika: III: Cichlidogyrus infecting the world's biggest cichlid and the non-endemic tribes Haplochromini, Oreochromini and Tylochromini (Teleostei, Cichlidae).

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.

Two new species of protogyrodactylus (Monogenoidea: Dactylogyridae) from the gills of Gerres nigri (Teleostei: Gerreidae) from Senegal.

Protogyrodactylus ethiopicus sp. n. and P. kritskyi sp. n. are described from the gills of Gerres nigri Günther (Gerreidae, Perciformes) captured from the estuary of the Sine-Saloum River (Senegal, West Africa). These new species differ from previously described species within the genus by a mid- or dextro-ventral vaginal opening (dextral in all other species). They are part of a morphological species group within Protogyrodactylus Johnston et Tiegs, 1922 that has the tip of the superficial root of the ventral anchor resembling a hook and two anterior projections on the anterior margin of the ventral bar. Protogyrodactylus ethiopicus differs from the remaining species in this group mainly by the morphology of the base of the male copulatory organ (MCO), which is disk-shaped, and the shape of the anterior projections of the ventral bar (round in the new species and relatively elongate in the other species of the group). The other new species, P. kritskyi, differs from all others in the same morphological group in having a MCO with a greatly expanded base that bears a heel-like subterminal sclerotization.

Two new Quadriacanthus Species (Monogenea: Dactylogyridae) Including a New Geographical Record of Quadriacanthus mandibulatus Francová & Rehulková, 2017, Gill Parasites from Heterobranchus bidorsalis (Siluriformes) in Côte d'Ivoire.

PURPOSE: To study the monogeneans gill parasites of Heterobranchus bidorsalis (Siluriformes, Clariidae) fish of economic interest in the Bagoue River. METHODS: Fifty-five specimens of H. bidorsalis were caught with gillnets in the Bagoué River from August 2018 to January 2019. The monogeneans were individually isolated from the gill and transferred directly onto a slide in a drop of glycerin ammonium picrate mixture (GAP). RESULTS: The new species differ from the previously described Quadriacanthus spp. mainly by the morphology of their sclerotised vagina: Quadriacanthus tanoyaoi sp. nov having a V-shaped, widely open vagina with a central protuberance, and Quadriacanthus bidorsalisi sp. nov having a pear-shaped vagina, composed of a tubular part with a ring near the aperture and a swollen portion. Q. tanoyaoi sp. nov and Q. bidorsalisi sp. nov are different in the shape of the dorsal and ventral anchors (more sturdy for Q. tanoyaoi), of the dorsal and ventral cuneus (triangular vs Y-shaped, respectively), and that of the vagina (V-shaped, widely open with a central protuberance vs pear-shaped, respectively). CONCLUSION: To date, only Quadriacanthus mandibulatus Francová & Rehulková, 2017 were recognised parasitizing from H. bidorsalis from the Nile River Basin (Sudan) and Lake Turkana (Kenya). The present study with the description of 2 new species expands the number to 3 species of monogeneans for this fish.