Taxonomic status of trichodina heterodentata Duncan, 1977 (Ciliophora: Peritrichia) using standard morphology, as well as molecular techniques

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Date
2017-12
Authors
De Jager, Gerhard P.
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University of the Free State
Abstract
English: The family Trichodinidae Raabe, 1959 (Ciliophora, Peritrichia) consists of 11 genera, of which the cosmopolitan genus Trichodina Ehrenberg, 1838 has the largest number of species. A substantial majority of trichodinid species are associated with marine and freshwater fish hosts, while some are also found associated with amphibians, as well as a variety of invertebrate hosts. Some trichodinid species show high levels of host specificity, whilst others occur on a range of hosts. Trichodina heterodentata Duncan, 1977 was first described from fish breeding farms in the Philippines as ectoparasites of imported cichlids, more specifically the Mozambique tilapia from southern Africa, Oreochromis mossambicus (Peters 1885), from the Limpopo River System. Trichodina heterodentata has subsequently been described from Europe, Australia, India, China, South America and Africa, however, it has not yet been found in North America. This cosmopolitan species, with a seeming preference for cichlid hosts, has unambiguous morphological features, but with distinct variances between and within populations. Most fish species harbouring T. heterodentata display multi-trichodinid infestations, however, on its amphibian host, T. heterodentata occurs as a single infestation, which makes it perfect for using it as a model species. After reviewing previous descriptions from North American trichodinids, analysing the morphological data (both from literature along with type material obtained from the Smithsonian Museum) and investigating the distribution patterns of the southern African introduced O. mossambicus throughout North American water systems, it seems likely that Trichodina hypsilepis Wellborn, 1967, is a synonym of T. heterodentata. This not only delegates several North American trichodinid species into synonymy, but also probably indicate an insidious African alien introduction sneaking it way into this continent. Morphological analysis of T. heterodentata, shows that this species is highly variable in regards to its denticle structure, which leads one to wonder if it’s might be a species complex, or indeed a single species. Trichodinid taxonomic work to date has primarily focused on morphological characteristics together with host and distribution records. With the advent of modern biochemical laboratory procedures, more and more emphasis is placed on molecular techniques in all fields of taxonomy, especially in parasitology. However, little molecular taxonomy work has been done on trichodinids. Trichodinid specimens were obtained from six different isolated host tadpole (Schlerophrys spp.) populations on the Nxamasere Flood plains, Okavango Panhandle, Botswana. This dissertation will represent the first study in southern Africa to successfully isolate genomic DNA from mobiline symbiotic ciliophorans. Genomic DNA was isolated from T. heterodentata and the 18S rDNA gene was amplified by PCR. After Sanger sequencing a complete phylogenetic tree of all available trichodinid sequences were constructed. All results indicate that trichodinids collected from tadpole hosts in the Okavango Delta are not T. heterodentata, but rather T. hypsilepis, supporting the theory that T. heterodentata, as it is known to date, is a species complex.
Afrikaans: Die familie Trichodinidae Raabe, 1959 (Ciliophora, Peritrichia) bestaan uit 11 genera, waarvan die mees kosmopolitiese genus, Trichodina Ehrenberg, 1838 die meeste spesies huisves. Die oorgrote meerderheid trigodina-spesies word met mariene en varswater visgashere geassosieër, terwyl ander op amfibiërs en ‘n verskeidenheid ander invertebrate gevind word. Sommige trigodinas is baie gasheerspesifiek en sommige word by talle verskillende tipe gashere aangetref. Trichodina heterodentata Duncan, 1977 was die eerste keer vanaf visteëlplase in die Filippyne op ingevoerde vis beskryf, meer spesifiek vanaf die ingevoerde bloukurpers,Oreochromis mossambicus (Peters 1885) uit die suider-Afrikaanse Limpopo Rivierstelsel. Trichodina heterodentata is sedertdien vanuit Europa, Australië, Indië, Sjina, Suid-Amerika en Afrika beskryf, maar nog nooit in Noord-Amerika nie. Hierdie kosmopolitiese spesie, met ‘n blykbare voorkeur vir Cichlidae gashere, het unieke morfologiese eienskappe, maar daar heers ook baie duidelike variasies tussen en binne verskillende bevolkings. Trichodina heterodentata word op meeste visspesies aangetref as deel van ‘n multi-spesie infestasie, alhoewel, op ‘n amfibiese gasheer blyk dit om ‘n enkel spesie infestasie te wees, wat hierdie gasheer ‘n ideale modelgasheer vir die studie van T. heterodentata maak. Na ‘n ondersoek van Noord-Amerikaanse trigodinas (vanaf literatuur en tiepmateriaal verkry vanaf die Smithsonianmuseum) en die verspreidingspatrone van O. mossambicus, blyk dit moontlik dat T. hypsilepis Wellborn, 1967 ‘n sinoniem van T. heterodentata kan wees. Laasgenoemde bevindings sink nie net sommige Noord-Amerikaanse trigodina spesies nie, maar verklaar ook hoe ‘n indringerspesie vannaf Afrika die pad na die Noord-Amerikaanse kontinent kon meemaak. As gevolg van die groot variasie wat T. heterodentata se tandringstrukture vertoon, word die vraag, of hierdie ‘n spesieskompleks of inderdaad ‘n enkel spesie is, gevra. Tot op hede het meeste van trigodina se taksonomie op morfologiese eienskappe en gasheerverspreiding berus. Selfs met die ontwikkeling van die moderne biochemiese laboratorium prosedure, waar al hoe meer gewig op molekulêre tegnieke (veral in parasitologie), maar ook in alle velde van taksonomie, geplaas word, is daar weinig molekulêre navorsing op trigodina-taksonomie gedoen. Trigodinas is vanaf ses verskillende paddavisgasheerbevolkings in die Nxamasere-vloedvlakte in die Okavango-pypsteel, Botswana versamel. Hierdie verhandeling sal die eerste studie in suiderlike-Afrika wees waar genomiese DNS vannaf simbiotiese siliate suksesvol geïsoleer word. Genomiese DNS is vanaf T. heterodentata geïsoleer en die 18S klein subeenheid rDNS deur PCR geamplifiseer. Na Sanger-verlenging is ‘n volledige filogenetiese boom saamgestel met al die beskikbare trigodina-gene. Die resultate dui aan dat die trigodinas vanaf paddavisse in die Okavango-delta nie T. heterodentata, soos aanvanklik gespekuleer, is nie, maar eerder T. hypsilepis. Dit ondersteun dus die teorie dat T. heterodentata, soos tans bekend is, ‘n spesies-kompleks is.
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Trichodina heterodenta, Trichodina hypsilepis, Oreochromis mossambicus, Schlerophrys gutteralis, Nxamasere floodplain, Okavango panhandle, Morphology, Molecular, Taxonomy, Schlerophrys poweri, Dissertation (M.Sc. (Zoology and Entomology))--University of the Free State, 2017
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