Revision of Nannocharax luapulae Boulenger, 1915 (Characiformes: Distichodontidae) from the Upper Congo basin: Evidence for a species pair.

For many decades, Nannocharax luapulae has been considered to be widespread in the southern part of the Upper Congo basin. However, meristic, morphometric and cytochrome c oxidase subunit I (COI) barcoding evidence revealed that its geographical distribution is restricted to the Luapula-Moero basin. The populations of the Upper Lualaba are assigned to a new species, N. chochamandai. This new species, though highly similar to N. luapulae, can readily be distinguished from it by its lower number of lateral line scales, 41-46 (vs. 49-55), its pectoral fin reaching the pelvic-fin insertion (vs. not reaching the pelvic-fin insertion) and its pelvic fin reaching the base of the anal fin (vs. not reaching the base of the anal fin). Specimens of N. chochamandai display thickened pads on the first three pelvic-fin rays that exhibit intraspecific variation in development, which appears to be related to the flow-strength of the river in which these Nannocharax specimens occur. Nannocharax luapulae is redescribed and an updated identification key to the Nannocharax species of the Congo basin sensu lato is provided as well. Some fish conservation issues related to N. luapulae and N. chochamandai are also highlighted.

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.

Population genomics of introduced Nile tilapia Oreochromis niloticus (Linnaeus, 1758) in the Democratic Republic of the Congo: Repeated introductions since colonial times with multiple sources.

During colonial times, Nile tilapia Oreochromis niloticus (Linnaeus, 1758) was introduced into non-native parts of the Congo Basin (Democratic Republic of the Congo, DRC) for the first time. Currently, it is the most farmed cichlid in the DRC, and is present throughout the Congo Basin. Although Nile tilapia has been reported as an invasive species, documentation of historical introductions into this basin and its consequences are scant. Here, we study the genetic consequences of these introductions by genotyping 213 Nile tilapia from native and introduced regions, focusing on the Congo Basin. Additionally, 48 specimens from 16 other tilapia species were included to test for hybridization. Using RAD sequencing (27,611 single nucleotide polymorphisms), we discovered genetic admixture with other tilapia species in several morphologically identified Nile tilapia from the Congo Basin, reflecting their ability to interbreed and the potential threat they pose to the genetic integrity of native tilapias. Nile tilapia populations from the Upper Congo and those from the Middle-Lower Congo are strongly differentiated. The former show genetic similarity to Nile tilapia from the White Nile, while specimens from the Benue Basin and Lake Kariba are similar to Nile tilapia from the Middle-Lower Congo, suggesting independent introductions using different sources. We conclude that the presence of Nile tilapia in the Congo Basin results from independent introductions, reflecting the dynamic aquaculture history, and that their introduction probably leads to genetic interactions with native tilapias, which could lower their fitness. We therefore urge avoiding further introductions of Nile tilapia in non-native regions and to use native tilapias in future aquaculture efforts.

Association between Labeobarbus spp. (Teleostei: Cyprinidae) and environmental variables in the Luhoho basin (Eastern Congo River basin; DRC).

The African cyprinid genus Labeobarbus is characterized by a high polymorphism in mouth morphology. The association between four species of this genus (two chiselmouths, one rubberlip with a well-developed fleshy mental lobe on the lower jaw, i.e., a fleshy appendage on the chin, and one usually with a poorly developed one) and 26 environmental variables was investigated in four affluents of the Luhoho River in the Albertine Highlands of the Upper Congo basin. A combination of univariate and multivariate analyses suggested that differences exist in habitat composition at localities where different Labeobarbus species were reported, and within different mouth phenotypes. The association of L. longidorsalis with the presence of cobbles and pebbles and of L. brauni with the presence of boulders suggests that the occurrence of chiselmouths phenotype is associated with stony habitats, whereas phenotypes with a mental lobe, i.e., Labeobarbus longifilis and L. paucisquamatus, mostly occupy habitats without stones. Differences in the distribution between these two species appear to be linked to physico-chemical parameters such as electrical conductivity and water temperature rather than substrate type. The findings of this study are relevant for developing guidelines that aim effective local preservation and conservation of these Labeobarbus species.

The synonymy of Haplochromis pharyngalis and Haplochromis petronius (Cichlidae).

Haplochromis pharyngalis and Haplochromis petronius, two endemic cichlids from the Lake Edward system (Uganda, Democratic Republic of the Congo), are very similar in general morphology but have been reported to differ in pharyngeal jaw morphology and distribution. This study analysed 51 morphometrics and various qualitative characteristics of 48 specimens from different localities. The morphological traits of both species strongly overlap, and differences in the pharyngeal jaw morphology correspond to a geographic morphocline. We conclude that all specimens belong to one valid species, H. pharyngalis, and consider H. petronius to be a synonym.

Mitogenomic characterization and systematic placement of the Congo blind barb Caecobarbus geertsii (Cypriniformes: Cyprinidae).

This study presents the first complete mitochondrial genome (mitogenome) of Caecobarbus geertsii, the Congo blind barb, a cave-dwelling, CITES-protected, cyprinid fish endemic to the Lower Congo basin (DRC). The length of the circular mitogenome is 16,565 base pairs. The 13 protein coding genes, 2 ribosomal RNA genes and 22 transfer RNA genes are similar in position and direction to those of other members of the family Cyprinidae. Phylogenetic analyses including 28 complete mitogenomes from representatives of the subfamily Smiliogastrinae (Cyprinidae), showed that Caecobarbus was nested within a clade including representatives of the genus Enteromius. The data presented in this study provide information on the molecular identification and classification of this threatened species. The results further suggest the need for a taxonomic revision of the genus Enteromius.

An annotated checklist of the fish fauna of the river systems draining the Kahuzi-Biega National Park (Upper Congo: Eastern DR Congo).

The Kahuzi-Biega National Park (KBNP), situated mainly in the Eastern Highlands Ecoregion of the Upper Congo basin, is drained by the Lowa and Ulindi rivers, and some western affluents of Lake Kivu. In this study, the first list of the fish diversity of these systems is provided based on museum collections and complemented, for the Lowa River system and the western Lake Kivu affluents, with recently collected specimens (2013-2017). A total of 118 species are reported from the Lowa basin, 22 from the Ulindi basin and seven from these Lake Kivu affluents. Within the Lowa and Ulindi, respectively, five and one species, all cichlids, have been introduced. Currently, 51 species are reported from within the park, only two of which have been reported from the highlands, i.e., Amphilius kivuensis from the Luha, the source of the Luka River, and Clarias liocephalus from the headwaters of the Lake Kivu' affluents. With a total of 30 species, Cyprinidae is by far the largest family, representing 25% of the total species diversity of the Lowa basin. It is followed by Mormyridae with 13 species (11%), Alestidae and Mochokidae with 10 species each (8%), Clariidae and Amphiliidae with eight species (7%), and Distichodontidae with six species (5%). Seven new species for science were discovered and 11 species were found to be endemic to the Lowa system. Although further exploration is needed, this underscores the importance of the KBNP in protecting the fish fauna of the Lowa basin but also highlights the park's limited coverage of the fish fauna of the Lowa basin.

Where ichthyofaunal provinces meet: the fish fauna of the Lake Edward system, East Africa.

Based on literature, museum collections and three recent expeditions, an annotated species list of the Lake Edward, East Africa, drainage system is presented, excluding the endemic haplochromines. A total of 34 non-Haplochromis species belonging to 10 families and 21 genera are recorded from the system. Three of these are endemic and two others have been introduced in the region. Six species are new records for the Lake Edward system. A species accumulation curve indicates that we probably covered most of the non-Haplochromis species in the area sampled during the recent expeditions, but undetected species might still be present in the Congolese part of the system, which is poorly sampled. A comparison of the species list with those of neighbouring basins confirmed the placement of the Lake Edward system within the east-coast ichthyofaunal province.

Similar ecology, different morphology: Three new species of oral-mollusc shellers from Lake Edward.

Lake Edward, East Africa, harbours a largely understudied assemblage of haplochromine cichlids that displays a range of adaptions to various specialised trophic niches. In this system, we discovered specimens of Haplochromis with morphologies similar to those of oral-mollusc shellers from Lake Victoria. These morphologies are characterised by short oral jaws with stout teeth that are used either to crush molluscs or to grab the soft bodies of snails and wrench them out of their shells. A morphometric study on 47 specimens from Lake Edward revealed the presence of three new species with an oral-shelling morphology: Haplochromis concilians sp. nov., H. erutus sp. nov. and H. planus sp. nov. All three species are formally described. Stomach-content observations confirmed an opportunistic oral-shelling ecology for H. concilians sp. nov. and H. erutus sp. nov. Within H. planus sp. nov., only large specimens displayed a specialised oral-shelling morphology, but their stomachs were nearly empty, while small specimens consumed mainly Ostracoda and Hydrachnidia. Remarkably, the three species differed considerably in morphology from each other, but they each resembled oral-sheller species from Lake Victoria.

Enteromius thespesios (Teleostei: Cyprinidae): a new minnow species with a remarkable sexual dimorphism from the south-eastern part of the Upper Congo River.

A new minnow species, Enteromius thespesios, is described from the south-eastern part of the upper Congo River; that is, the Kalule Nord, the Luvilombo and the Chambeshi Rivers. Enteromius thespesios belongs to the group of the soft-rayed species of Enteromius from the Congo Basin; that is, those with a weakly ossified, flexible last unbranched dorsal-fin ray that lacks serrations along its posterior edge. Within this group, E. thespesios is most similar to E. humeralis, from which it is distinguished by a higher number of circumpeduncular scales and shorter anterior and posterior barbels. Enteromius thespesios is a rheophilic and territorial species. It exhibits a marked sexual dimorphism, with males having: a red band towards the distal edge of dorsal, caudal and, to a lesser degree, anal fin; nuptial tubercles; a longer snout; longer pectoral fins; a shorter anal fin. This study gives extensive consideration to sexual shape differences for a species of Enteromius and also briefly reviews the current knowledge of sexual dimorphism in the species of Enteromius from the Congo Basin. Some conservation issues related to the new species are also highlighted.

Terrestrial contributions to Afrotropical aquatic food webs: The Congo River case.

Understanding the degree to which aquatic and terrestrial primary production fuel tropical aquatic food webs remains poorly understood, and quantifying the relative contributions of autochthonous and allochthonous inputs is methodologically challenging. Carbon and nitrogen stable isotope ratios (δ 13C, δ 15N) can provide valuable insights about contributions of terrestrial resources and trophic position, respectively, but this approach has caveats when applied in typical complex natural food webs.Here, we used a combination of C, N, and H (δ 2H) stable isotope measurements and Bayesian mixing models to estimate the contribution of terrestrial (allochthonous) and aquatic (autochthonous) inputs to fish and invertebrate communities in the Congo River (and some tributaries).Overall, our results show that we gained power to distinguish sources by using a multiple tracer approach and we were able to discriminate aquatic versus terrestrial sources (esp. including hydrogen isotopes). Fish δ 2H values were clearly correlated with their food preferences and revealed a high level of variation in the degree of allochthony in these tropical aquatic communities.At the community level, it is clear that terrestrial C3 plants are an important source fueling the Congo River food web. However, in order to better constrain source contribution in these complex environments will require more robust constraints on stable isotope values of algal and methane-derived C sources.

DNA barcoding fishes from the Congo and the Lower Guinean provinces: Assembling a reference library for poorly inventoried fauna.

The Congolese and Lower Guinean ichthyological provinces are understudied hotspots of the global fish diversity. Here, we barcoded 741 specimens from the Lower and Middle Congo River and from three major drainage basins of the Lower Guinean ichthyological province, Kouilou-Niari, Nyanga and Ogowe. We identified 194 morphospecies belonging to 82 genera and 25 families. Most morphospecies (92.8%) corresponded to distinct clusters of DNA barcodes. Of the four morphospecies present in both neighbouring ichthyological provinces, only one showed DNA barcode divergence <2.5%. A small fraction of the fishes barcoded here (12.9% of the morphospecies and 16.1% of the barcode clusters representing putative species) were also barcoded in a previous large-scale DNA analysis of freshwater fishes of the Lower Congo published in 2011 (191 specimens, 102 morphospecies). We compared species assignments before and after taxonomic updates and across studies performed by independent research teams and observed that most cases of inconsistent species assignments were due to unknown diversity (undescribed species and unknown intraspecific variation). Our results report more than 17 putative new species and show that DNA barcode data provide a measure of genetic variability that facilitates the inventory of underexplored ichthyofaunae. However, taxonomic scrutiny, associated with revisions and new species descriptions, is indispensable to delimit species and build a coherent reference library.

Six new dactylogyrid species (Platyhelminthes, Monogenea) from the gills of cichlids (Teleostei, Cichliformes) from the Lower Congo Basin.

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.

Delineating species along shifting shorelines: Tropheus (Teleostei, Cichlidae) from the southern subbasin of Lake Tanganyika.

BACKGROUND: Species delineation is particularly challenging in taxa with substantial intra-specific variation. In systematic studies of fishes, meristics and linear measurements that describe shape are often used to delineate species. Yet, little is known about the taxonomic value of these two types of morphological characteristics. Here, we used Tropheus (Teleostei, Cichlidae) from the southern subbasin of Lake Tanganyika to test which of these types of characters best matched genetic lineages that could represent species in this group of stenotypic rock-dwelling cichlids. We further investigated intra-population variation in morphology. By linking this to a proxy of a population's age, we could assess the evolutionary stability of different kinds of morphological markers. RESULTS: Morphological data was collected from 570 specimens originating from 86 localities. An AFLP approach revealed the presence of five lineages in the southern subbasin: T. moorii, T. brichardi, T. sp. 'maculatus', T. sp. 'Mpimbwe' and T. sp. 'red', which we consider to represent distinct species. Although both types of morphological data supported this classification, a comparison of PST-values that describe inter-population morphological differentiation, revealed a better correspondence between the taxon delineation based on AFLP data and the patterns revealed by an analysis of meristics than between the AFLP-based taxon delineation and the patterns revealed by an analysis of shape. However, classifying southern populations of Tropheus was inherently difficult as they contained a large amount of clinal variation, both in genetic and in morphological data, and both within and among species. A scenario is put forward to explain the current-day distribution of the species and colour varieties and the observed clinal variation across the subbasin's shoreline. Additionally, we observed that variation in shape was larger in populations from shallow shores whereas populations from steep shores were more variable in meristics. This difference is explained in terms of the different timescales at which small and large scale lake level fluctuations affected populations of littoral cichlids at steep and shallow shores. CONCLUSIONS: Our results showed meristics to be more evolutionary stable, and of higher taxonomic value for species delimitation in Tropheus, than linear measurements that describe shape. These results should be taken into account when interpreting morphological differences between populations of highly stenotypic species, such as littoral cichlids from the Great East African Lakes.

Trematocranus pachychilus, a new endemic cichlid from Lake Malawi (Teleostei, Cichlidae).

A new species of Trematocranus, T. pachychilussp. n., is described from Lake Malawi. So far, it has only been found at Jafua Bay, Mozambique. It can easily be distinguished from T. labifer by its molariform pharyngeal dentition. A morphometric study, including 24 measurements and 15 counts, was done to compare the new species with T. microstoma and T. placodon. Trematocranus pachychilus is characterised by its thick lips. This species further differs from T. microstoma by its bicuspid (vs. unicuspid) outer oral teeth, wide (vs. small) pharyngeal bone, and its head shape. It resembles T. placodon, from which it can be distinguished by its straight to concave head profile (vs. rounded), less-developed pharyngeal bones (vs. hypertrophied), and the presence of small to minute teeth on the lateral parts of the dentigerous area on the lower pharyngeal bone. A key to the species of Trematocranus is provided.

Limited possibilities for prezygotic barriers in the reproductive behaviour of sympatric Ophthalmotilapia species (Teleostei, Cichlidae).

Since prezygotic rather than postzygotic barriers are believed to maintain the diversity of closely related sympatric cichlids, differences in phenotypic traits and reproductive behaviours are likely involved in maintaining species boundaries. Here, we focused on the reproductive behaviour of three Ophthalmotilapia species with distributions that only overlap on a small stretch of the shore line of Lake Tanganyika. Repeated introgression of mitochondrial DNA between these species was previously reported, which suggested they can hybridise. Our aim is to test the hypothesis that reproductive behaviour acts as a prezygotic barrier that prevents frequent hybridisation in sympatric Ophthalmotilapia species. We performed a quantitative analysis of twelve reproductions (four for O. ventralis, six for O. nasuta, one for O. boops, and one between a female O. ventralis and a male O. nasuta). Although similar ethograms were obtained for these reproductions, the O. ventralis and O. boops males displayed a behaviour that was never performed by O. nasuta males. This behaviour was displayed during courtship and we called it 'invite'. In O. ventralis, we could show that it was associated with the emission of a single pulse sound. The comparison of O. nasuta and O. ventralis reproductive behaviours also revealed some quantitative differences: O. ventralis males showed the location of the bower more often to the female, whereas O. ventralis females followed the male more often. The similarity between the reproductive behaviours in O. ventralis and O. nasuta could explain the occurrence of the heterospecific spawning event recorded between an O. nasuta male and an O. ventralis female. Importantly, few eggs were laid and the maternal mouthbrooding that resulted from this heterospecific reproduction only lasted for two days, which suggested the abortion of egg development. Hence, in the absence of conspecifics, courtship and mating behaviours alone do not constitute perfect prezygotic barriers between these two species.

A new species of Pseudocrenilabrus (Perciformes: Cichlidae) from Lake Mweru in the Upper Congo River System.

Pseudocrenilabrus pyrrhocaudalis sp. nov. is described from Lake Mweru in the upper Congo River drainage, on the border of the Democratic Republic of Congo and Zambia. This species, which appears to be endemic to the lake, lives in sympatry with P. philander. Pseudocrenilabrus pyrrhocaudalis sp. nov. is distinguished from P. philander in nuptial males by the presence of an orange colour on the ventral part of the body and the proximal parts of the anal and caudal fins, a broad band of bright white on the distal edge of anal and caudal fins, a uniform grey head and dorsum, and a subtruncate caudal fin. In addition, P. pyrrhocaudalis has a shorter snout, a narrower head, a smaller interorbital distance, a smaller pre-anal distance, a more slender caudal peduncle and fewer scales around the caudal peduncle in both sexes.

Testing the potential of DNA barcoding in vertebrate radiations: the case of the littoral cichlids (Pisces, Perciformes, Cichlidae) from Lake Tanganyika.

We obtained 398 cytochrome c oxidase subunit I barcodes of 96 morphospecies of Lake Tanganyika (LT) cichlids from the littoral zone. The potential of DNA barcoding in these fishes was tested using both species identification and species delineation methods. The best match (BM) and best close match (BCM) methods were used to evaluate the overall identification success. For this, three libraries were analysed in which the specimens were categorized into Operational Taxonomic Units (OTU) in three alternative ways: (A) morphologically distinct, including undescribed, species, (B) valid species and (C) complexes of morphologically similar or closely related species. For libraries A, B and C, 73, 73 and 96% (BM) and 72, 70 and 94% (BCM) of the specimens were correctly identified. Additionally, the potential of two species delineation methods was tested. The General Mixed Yule Coalescent (GMYC) analysis suggested 70 hypothetical species, while the Automatic Barcode Gap Discovery (ABGD) method revealed 115 putative species. Although the ABGD method had a tendency to oversplit, it outperformed the GMYC analysis in retrieving the species. In most cases where ABGD suggested oversplitting, this was due to intraspecific geographical variation. The failure of the GMYC method to retrieve many species could be attributed to discrepancies between mitochondrial gene trees and the evolutionary histories of LT cichlid species. Littoral LT cichlids have complex evolutionary histories that include instances of hybridization, introgression and rapid speciation. Nevertheless, although the utility of DNA barcoding in identification is restricted to the level of complexes, it has potential for species discovery in cichlid radiations.

Taxonomic challenges in freshwater fishes: a mismatch between morphology and DNA barcoding in fish of the north-eastern part of the Congo basin.

This study evaluates the utility of DNA barcoding to traditional morphology-based species identifications for the fish fauna of the north-eastern Congo basin. We compared DNA sequences (COI) of 821 samples from 206 morphologically identified species. Best match, best close match and all species barcoding analyses resulted in a rather low identification success of 87.5%, 84.5% and 64.1%, respectively. The ratio 'nearest-neighbour distance/maximum intraspecific divergence' was lower than 1 for 26.1% of the samples, indicating possible taxonomic problems. In ten genera, belonging to six families, the number of species inferred from mtDNA data exceeded the number of species identified using morphological features; and in four cases indications of possible synonymy were detected. Finally, the DNA barcodes confirmed previously known identification problems within certain genera of the Clariidae, Cyprinidae and Mormyridae. Our results underscore the large number of taxonomic problems lingering in the taxonomy of the fish fauna of the Congo basin and illustrate why DNA barcodes will contribute to future efforts to compile a reliable taxonomic inventory of the Congo basin fish fauna. Therefore, the obtained barcodes were deposited in the reference barcode library of the Barcode of Life Initiative.

Hidden biodiversity in an ancient lake: phylogenetic congruence between Lake Tanganyika tropheine cichlids and their monogenean flatworm parasites.

The stunning diversity of cichlid fishes has greatly enhanced our understanding of speciation and radiation. Little is known about the evolution of cichlid parasites. Parasites are abundant components of biodiversity, whose diversity typically exceeds that of their hosts. In the first comprehensive phylogenetic parasitological analysis of a vertebrate radiation, we study monogenean parasites infecting tropheine cichlids from Lake Tanganyika. Monogeneans are flatworms usually infecting the body surface and gills of fishes. In contrast to many other parasites, they depend only on a single host species to complete their lifecycle. Our spatially comprehensive combined nuclear-mitochondrial DNA dataset of the parasites covering almost all tropheine host species (N = 18), reveals species-rich parasite assemblages and shows consistent host-specificity. Statistical comparisons of host and parasite phylogenies based on distance and topology-based tests demonstrate significant congruence and suggest that host-switching is rare. Molecular rate evaluation indicates that species of Cichlidogyrus probably diverged synchronically with the initial radiation of the tropheines. They further diversified through within-host speciation into an overlooked species radiation. The unique life history and specialisation of certain parasite groups has profound evolutionary consequences. Hence, evolutionary parasitology adds a new dimension to the study of biodiversity hotspots like Lake Tanganyika.