Search

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., (© T. Moons et al., published by EDP Sciences, 2023.) more...

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., (© 2022 Willi Hennig Society.) more...

We describe a new genus of dalytyphloplanid rhabdocoels, with seven new species. Orostylis gen. nov. has a unique combination of characters including the presence of a sclerotised stylet, an anteriorly positioned male copulatory organ with the male genital pore in the buccal cavity, and the absence of an oviduct. The ovary empties directly into the intestinal lumen. Orostylis dohae sp. nov., Orostylis asinaraensis sp. nov., Orostylis caecus sp. nov., Orostylis distortus sp. nov., Orostylis donanae sp. nov., Orostylis gallicus sp. nov., and Orostylis timucuorum sp. nov. are distinguished from each other by the structure of the sclerotised parts of the male copulatory organ. Molecular and morphological evidence place the new genus in Neodalyellida Willems et al. 2006, and similarities with other taxa are discussed. more...

A comprehensive morphological and taxonomic account of the members of the genus Cheliplana de Beauchamp, 1927 is presented. Six new species are described: Cheliplana asinaraensis n. sp., C. cubana n. sp., C. curacaoensis n. sp., C. hawaiiensis n. sp., C. longissima n. sp. and C. mauii n. sp. The new species are mainly distinguished from each other and from other representatives of Cheliplana by the organisation of the reproductive system and the structure of the cirrus. Furthermore, C. triductibus Van Steenkiste, Volonterio, Schockaert Artois, 2008 is considered a junior synonym of Cheliplana deverticula Ax, 2008. The two subspecies of Cheliplana asica Marcus, 1952, C. asica asica and C. asica terminalis Brunet, 1968, are considered separate species. The systematic position of the genus Dactyloplana Armonies, 2018 is discussed, and its synonymy with Cheliplana is retained. As such, this brings the total number of species of Cheliplana to 49. Finally, we provide an identification key to the members of the genus, based on characters that enable identification to species level in the field. more...

The taxon Koinocystididae is the third most species-rich family within Eukalyptorhynchia. However, its diversity and phylogeny have been largely neglected in former studies. We introduce three new genera and twelve new species of Koinocystididae including Simplexcystis asymmetrica gen. n. sp. n., Galapagetula cubensis sp. n., eight species of Reinhardorhynchus gen. n. and two species of Itaipusa. This raises the total number of species within Koinocystididae from 51 to 63. We also report on new distribution records for six known species: I. divae (Cuba, Panama and New Caledonia), I. karlingi (Sardinia and Lanzarote), Reinhardorhynchus riegeri comb. n. (Cuba), R. ruffinjonesi comb. n. (Cuba and Panama), Utelga heinckei (Cuba and Lanzarote), and U. pseudoheinckei (Sardinia). Simplexcystis asymmetrica gen. n. sp. n. is characterised by a male duct running eccentrically through the copulatory bulb, lack of any hard structures in the male system, lack of a bursa, and the fact that the epithelia of the female, the male, and part of the common atrium are covered by a brush border. Galapagetula cubensis sp. n. has a caudal gonopore, a divisa-type copulatory bulb with an unarmed penis papilla, and a female duct without a sphincter. The new species of Itaipusa and Reinhardorhynchus gen. n. differ from their congeners in the detailed structure of the copulatory bulb and especially the hard structures associated with it. In a molecular phylogenetic analysis based on all available 18S and 28S rDNA sequences of koinocystidids, we found support for the monophyly of the family and the genus Utelga Marcus, 1949. The genus Itaipusa is not monophyletic in that I. sinensis forms a clade with Rhinolasius dillonicus, while other species of Itaipusa that have a copulatory bulb armed with hooks form a clade together with Sekerana stolzi. As the type species of Itaipusa (I. divae) is in neither of these clades, we erected a new genus for I. sinensis (Koinogladius gen. n.) and one for species of Itaipusa having a hook-bearing copulatory bulb (Reinhardorhynchus gen. n.), respectively. Whether the remaining species of Itaipusa form a monophylum remains uncertain. more...

The first three mitochondrial (mt) genomes of endosymbiotic turbellarian flatworms are characterised for the rhabdocoels Graffilla buccinicola, Syndesmis echinorum and S. kurakaikina. Interspecific comparison of the three newly obtained sequences and the only previously characterised rhabdocoel, the free-living species Bothromesostoma personatum, reveals high mt genomic variability, including numerous rearrangements. The first intrageneric comparison within rhabdocoels shows that gene order is not fully conserved even between congeneric species. Atp8, until recently assumed absent in flatworms, was putatively annotated in two sequences. Selection pressure was tested in a phylogenetic framework and is shown to be significantly relaxed in this and another protein-coding gene: cox1. If present, atp8 appears highly derived in platyhelminths and its functionality needs to be addressed in future research. Our findings for the first time allude to a large degree of undiscovered (mt) genomic plasticity in rhabdocoels. It merits further attention whether this variation is correlated with a symbiotic lifestyle. Our results illustrate that this phenomenon is widespread in flatworms as a whole and not exclusive to the better-studied neodermatans., Competing Interests: Declaration of competing interest None., (Crown Copyright © 2020. Published by Elsevier B.V. All rights reserved.) more...

Lake Tanganyika, East Africa, is the oldest and deepest African Great Lake and harbours one of the most diverse fish assemblages on earth. Two clupeid fishes, Limnothrissa miodon and Stolothrissa tanganicae, constitute a major part of the total fish catch, making them indispensable for local food security. Parasites have been proposed as indicators of stock structure in highly mobile pelagic hosts. We examined the monogeneans Kapentagyrus limnotrissae and Kapentagyrus tanganicanus (Dactylogyridae) infecting these clupeids to explore the parasites' lake-wide population structure and patterns of demographic history. Samples were collected at seven sites distributed across three sub-basins of the lake. Intraspecific morphological variation of the monogeneans (n = 380) was analysed using morphometrics and geomorphometrics of sclerotised structures. Genetic population structure of both parasite species (n = 246) was assessed based on a 415 bp fragment of the mitochondrial cytochrome c oxidase subunit I (COI) gene. Overall, we observed a lack of clear geographical morphological differentiation in both parasites along a north-south axis. This lack of geographical population structure was also reflected by a large proportion of shared haplotypes, and a pattern of seemingly unrestricted gene flow between populations. Significant morphological and genetic differentiation between some populations might reflect temporal differentiation rather than geographical isolation. Overall, the shallow population structure of both species of Kapentagyrus reflects the near-panmictic population structure of both host species as previously reported. Morphological differences related to host species identity of K. tanganicanus were consistent with incipient speciation at the genetic level. Both parasite species experienced a recent demographic expansion, which might be linked to paleohydrological events. Finally, interspecific hybridisation was found in Kapentagyrus, representing the first case in dactylogyrid monogeneans., (Copyright © 2020 Australian Society for Parasitology. Published by Elsevier Ltd. All rights reserved.) more...

Coelogynopora schockaerti n. sp. (Proseriata: Coelogynoporidae) is described from the northwestern Gulf of Lion, Mediterranean. Only a few representatives of this genus are reported from Mediterranean marine sandy shores and only from the northwestern to central Mediterranean, but all of these species also occur outside the Mediterranean. So far, C. schockaerti n. sp. is the only species exclusively known from the Mediterranean. The species is solely encountered in the Gulf of Lion, a coldwater pocket in the Mediterranean and can, therefore, be considered as a remnant of an ancient coldwater platyhelminth fauna. C. schockaerti n. sp. is characterized by the shape and size of the spines of the male copulatory organ and the presence, orientation, shape and size of the two accessory spines.  An identification key is given for the entire genus. more...

We report on the schizorhynch species collected in a survey in the eastern region of Cuba. Eighteen species were identified, of which only three are known to science: Cheliplana asica, C. terminalis, and Carcharodorhynchus flavidus. The 15 new species belong to three different genera: Cheliplana (five species), Carcharodorhynchus (four species), and Schizochilus (six species). The five new species of Cheliplana (C. gibarenha sp. n., C. santiaguera sp. n., C. spuriaseminalis sp. n., C. subproximalis sp. n. and C. verrucosa sp. n.) differ from their congeners in the detailed morphology of the proboscis hooks and the atrial organs. The proboscis hooks bear a distal small hook in C. verrucosa sp. n., a feature only shared with C. paradoxa. Two of the four new species of Carcharodorhynchus (C. smilodon sp. n. and C. papillaris sp. n. ) are very similar to C. flavidus in that the toothed belts of the proboscis are not continuous. However, they clearly differ from that species and from each other in the detailed construction of the teeth and copulatory organ. Carcharodorhynchus spiniformis sp. n. and C. nativus sp. n. can also be distinguished from the other species of Carcharodorhynchus by the detailed construction of the proboscis teeth and copulatory organ. The six new species of Schizochilus here described show a spiny cirrus around a central stylet. Two species lack the distal sclerotized cap of the copulatory bulb: S. favus sp. n. and S. bueycabonensis sp. n.. All new species of Schizochilus can be distinguished from each other and their congeners by the detailed construction of the hard parts (cirrus and stylet) of the copulatory organ. more...

The first records of Polycystididae from Cuba are provided and discussed. In total nine species have been collected, five of which are new to science, one representing a new genus. Polycydora intermedia gen. n. sp. n. shows intermediate features between Polycystis Kölliker, 1845 and Paulodora Marcus, 1948. The ovaries are kidney shaped, with the oocytes arranged in a row, and lack the hard "nozzles" of Paulodora. A female bursa as in Polycystis is present. In P. intermedia gen. n. sp. n. lacks the strong muscle bulb at the male bursa and the accessory glands type I in the male atrium, which are present in Polycystis. The male atrial organs include a prostate stylet type II connected to a free prostate vesicle type II. The four new species (Phonorhynchoides minor sp. n., Phonorhynchopsis capillaris sp. n., Phonorhynchopsis sublinguatus sp. n., Myobulla armenterosi sp. n.) are distinguished from their congeners by the specific shape and length of the male hard copulatory structures. Phonorhynchoides minor sp. n. has the smallest stylets within the genus, and these stylets are more straight than those of the other species of the genus. In Phonorhynchopsis capillaris sp. n. the prostate stylet type IV is only 20% of the length of the accessory stylet type IV, the lowest relative length within the genus. The accessory stylet in this species is extremely thin; it is only 2 µm wide. Phonorhynchopsis sublinguatus sp. n. has a prostate stylet widened in the distal half, ending in a rounded tip, not twisted in the middle. Because of its strong resemblance to M. armenterosi sp. n., M. berti sp. n., a new species of Myobulla Artois Schockaert, 2000 collected at the Atlantic coast of Panama is described here too. Both species of Myobulla have a prostate stylet type III that shows a 90° turn at some point, as is the case in some other species of Myobulla. In M. berti sp. n. the stylet is smaller overall and is bent in the middle; in M. armenterosi sp. n. the stylet is larger and it is bent more distally. All of the four known species have a very wide geographical distribution: Phonorhynchopsis haegheni, Alcha evelinae, Paraustrorhynchus elixus, and Gyratrix hermaphroditus. more...

This study represents the first exploration of the parasite fauna of cichlid fishes in the Mweru-Luapula subregion (Central Africa). Twelve species of cichlids and 14 species of Monogenea from three genera (Cichlidogyrus, Gyrodactylus and Scutogyrus) were collected. We present a first record of the gill parasite fauna of eight host species, Oreochromis mweruensis, Orthochromis sp. 'Mambilima', Sargochromis mellandi, Serranochromis angusticeps, S. stappersi, S. thumbergi and Tylochromis mylodon. The host range of ten parasite species was expanded. The study further includes the description of Cichlidogyrus consobrini sp. n. from S. mellandi and Orthochromis sp. 'Mambilima'. A new morphotype of C. halli is characterized, and three species - C. papernastrema, C. quaestio and C. zambezensis - are redescribed. Furthermore, the biodiversity and host specificity of these parasites is compared with that of cichlid parasites from Lake Kariba and Cameroon. Two species, including C. consobrini sp. n. and a new morphotype of C. halli, are putative endemics. The parasite fauna in Bangweulu-Mweru is highly similar in species composition to Lake Kariba, but in Bangweulu-Mweru the same parasite species are more host-specific, probably because of hydrogeographical differences between the two regions. more...

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., (© M.W.P. Jorissen et al., published by EDP Sciences, 2018.) more...

One of the major challenges in the development of alternative carcinogenicity assays is the prediction of non-genotoxic carcinogens. The variety of non-genotoxic cancer pathways complicates the search for reliable parameters expressing their carcinogenicity. As non-genotoxic and genotoxic carcinogens have different cancer risks, the objective of this study was to develop a concept for an in vivo test, based on flatworm stem cell dynamics, to detect and classify carcinogenic compounds. Our methodology entails an exposure to carcinogenic compounds during the animal's regeneration process, which revealed differences in proliferative responses between non-genotoxic and genotoxic carcinogens during the initial stages of the regeneration process. A proof of concept was obtained after an extensive study of proliferation dynamics of a genotoxic and a non-genotoxic compound. A pilot validation with a limited set of compounds showed that the proposed concept not only enabled a simple prediction of genotoxic and non-genotoxic carcinogens, but also had the power to discriminate between both. We further optimized this discrimination by combining stem cell proliferation responses with a phenotypic screening and by using specific knockdowns. In the future, more compounds will be tested to further validate and prove this concept. more...

Predatory flatworms belonging to the taxon Kalyptorhynchia are characterized by an anterior muscular proboscis that they use to seize prey. In many cases, the proboscis is armed with hooks, derived either from the extracellular matrix that surrounds the muscles or from intracellular deposits in the epithelium covering the proboscis. Glands associated with the proboscis reportedly are venomous; however, there are few direct tests of this hypothesis. This article reviews the structure and current knowledge of the function of the proboscis in the Kalyptorhynchia, points to areas in which the current understanding of phylogenetic relationships within this taxon is incongruent with our hypothesis of how the proboscis evolved, and addresses areas in need of further research, especially as regards functional morphology and biomechanics., (© The Author 2015. Published by Oxford University Press on behalf of the Society for Integrative and Comparative Biology. All rights reserved. For permissions please email: journals.permissions@oup.com.) more...

In this study we elaborate the phylogeny of Dalytyphloplanida based on complete 18S rDNA (156 sequences) and partial 28S rDNA (125 sequences), using a Maximum Likelihood and a Bayesian Inference approach, in order to investigate the origin of a limnic or limnoterrestrial and of a symbiotic lifestyle in this large group of rhabditophoran flatworms. The results of our phylogenetic analyses and ancestral state reconstructions indicate that dalytyphloplanids have their origin in the marine environment and that there was one highly successful invasion of the freshwater environment, leading to a large radiation of limnic and limnoterrestrial dalytyphloplanids. This monophyletic freshwater clade, Limnotyphloplanida, comprises the taxa Dalyelliidae, Temnocephalida, and most Typhloplanidae. Temnocephalida can be considered ectosymbiotic Dalyelliidae as they are embedded within this group. Secondary returns to brackish water and marine environments occurred relatively frequently in several dalyeliid and typhloplanid taxa. Our phylogenies also show that, apart from the Limnotyphloplanida, there have been only few independent invasions of the limnic environment, and apparently these were not followed by spectacular speciation events. The distinct phylogenetic positions of the symbiotic taxa also suggest multiple origins of commensal and parasitic life strategies within Dalytyphloplanida. The previously established higher-level dalytyphloplanid clades are confirmed in our topologies, but many of the traditional families are not monophyletic. Alternative hypothesis testing constraining the monophyly of these families in the topologies and using the approximately unbiased test, also statistically rejects their monophyly. more...

Three new species of Polycystididae are described: Brachyrhynchus triplostylis n. gen. n. sp., B. acutus n. sp. and B. oosterlyncki n. sp. They are the first species of Polycystididae to be described with a second, single-walled accessory stylet connected to a small glandular vesicle in the male system. The three species can be distinguished from each other by the detailed morphology and the relative lengths of the three stylets in the male system. The relationships of these species with other representatives of Polycystididae are discussed. more...

The flatworm species Schmidtea mediterranea and Macrostomum lignano have become new and innovative model organisms in stem cell, regeneration and tissue homeostasis research. Because of their unique stem cell system, (lab) technical advantages and their phylogenetic position within the Metazoa, they are also ideal candidate model organisms for toxicity assays. As stress and biomarker screenings are often performed at the transcriptional level, the aim of this study was to establish a set of reference genes for qPCR experiments for these two model organisms in different stress situations. We examined the transcriptional stability of nine potential reference genes (actb, tubb, ck2, cox4, cys, rpl13, gapdh, gm2ap, plscr1) to assess those that are most stable during altered stress conditions (exposure to carcinogenic metals and salinity stress). The gene expression stability was evaluated by means of geNorm and NormFinder algorithms. Sets of best reference genes in these analyses varied between different stress situations, although gm2ap and actb were stably transcribed during all tested combinations. In order to demonstrate the impact of bad normalisation, the stress-specific gene hsp90 was normalised to different sets of reference genes. In contrast to the normalisation according to GeNorm and NormFinder, normalisation of hsp90 in Macrostomum lignano during cadmium stress did not show a significant difference when normalised to only gapdh. On the other hand an increase of variability was noticed when normalised to all nine tested reference genes together. Testing appropriate reference genes is therefore strongly advisable in every new experimental condition. more...

Spermiogenesis in Macrostomum lignano (Macrostomorpha, Rhabditophora) is described using light- and electron microscopy of the successive stages in sperm development. Ovoid spermatids develop to highly complex, elongated sperm possessing an undulating distal (anterior) process (or "feeler"), bristles, and a proximal (posterior) brush. In particular, we present a detailed account of the morphology and ontogeny of the bristles, describing for the first time the formation of a highly specialized bristle complex consisting of several parts. This complex is ultimately reduced when sperm are mature. The implications of the development of this bristle complex on both sperm maturation and the evolution and function of the bristles are discussed. The assumed homology between bristles and flagellae questioned., ((c) 2008 Wiley-Liss, Inc.) more...

Four new species of freshwater rhabdocoel flatworms from ephemeral rock pools in southeastern Botswana are described and discussed. Two of them, Syringoplana kolasai n. gen. n. sp. and Mesostoma thamagai n. sp. belong to the Typhloplanidae Graff, 1905. The unique construction of the excretory system is the main characteristic of S. kolasai. M. thamagai can be separated from other Mesostoma Ehrenberg, 1837 species by the presence of a bundle of eosinophilic glands at the transition from oviduct to seminal receptacle. The other two taxa, Gieysztoria isoldeae n. sp. and G. faubeli n. sp. belong to the Dalyelliidae Graff, 1905. G. isoldeae is characterised by the presence of four separate hollow spines in the male atrium, which are connected to two accessory glandular organs. G. faubeli can be separated from other Gieysztoria Ruebush and Hayes, 1939 species by the detailed construction of the stylet. Apart from these two species the occurrence of an unidentified Microdalyellia Gieysztor, 1938 species is mentioned. more...

A new species of polyclad flatworm from Papua New Guinea is described. It is found symbiotic in the ophiuroid Ophiothrix purpurea von Martens, 1867 (Echinodermata: Ophiuroidea). Apparently it belongs to the taxon Discoplana Bock, 1913 and can be distinguished from the six previously described Discoplana species by its very short ejaculatory duct and a penial papilla covered with a penial sheath, but without any true sclerotised structures such as a stylet or spines. The cladistic analysis of the Discoplana/ Euplana species, based on morphological features and including two outgroups, reveals that all species of Discoplana, except D. pacificola, form a monophyletic taxon, that is not a synonym of Euplana Girard, 1893. Therefore the name Discoplana is conserved and the new species will be described as Discoplana malagasensis sp. nov. A key for the Discoplana/Euplana group is provided. In this key the biogeographical distribution and possible synonyms are given. more...