Morphology and Molecular Phylogeny of Two Little-Known Species of Loxodes, L. kahli Dragesco & Njiné, 1971 and L. rostrum Müller, 1786 (Protist, Ciliophora, Karyorelictea)
Tóm tắt
The morphology and phylogeny of two little-known species, Loxodes kahli Dragesco & Njiné, 1971 and L. rostrumMüller, 1786, isolated from freshwater muddy sediments in China, were investigated based on live features, infraciliature, and small subunit ribosomal DNA (SSU rDNA) sequence data. Loxodes kahli is distinguished from its congeners mainly by the number and arrangement of macronuclei (6–17 in one row) and the number of right somatic ciliary rows (11–26). The Chinese populations of L. kahli also exhibit differences with other populations in terms of the body size and the number of right ciliary rows. The characteristics of L. rostrum are consistent with those of previous studies except for the number of right ciliary rows (9–10). The studied species were redefined based on the new information and previous descriptions. This study also gave a brief morphological summary of the species in the genus Loxodes by an identification key. SSU rDNA sequence-based phylogenetic analyses revealed that both species are grouped with their congeners, supporting the monophyly of the genus Loxodes.
Tài liệu tham khảo
Bobyleva, N. N., Kudrjavtsev, B. N., and Raikov, I. B., 1980. Changes of the DNA content of differentiating and adult macronuclei of the ciliate Loxodes magnus (Karyorelictida). Journal of Cell Science, 44 (8): 375.
Buonanno, F., 2005. Variations in the efficiency of ciliate extrusomal toxins against a common ciliate predator, the catenulid Stenostomum sphagnetorum. Italian Journal of Zoology, 72 (4): 293–295.
Buonanno, F., Saltalamacchia, P., and Miyake, A., 2005. Defence function of pigmentocysts in the karyorelictid ciliate Loxodes striatus. European Journal of Protistology, 41 (2): 151–158.
Dragesco, J., 1970. Ciliés libres du Cameroun. Annales de la Faculté des Sciences Université Fédérale du Cameroun (Numéro Hors-série). Université Fédérale du Cameroun, Yaoundé, 1–141 (in French).
Dragesco, J., and Njiné, T., 1971. Compléments à la connaissance des ciliés libres du Cameroun. Annales de la Faculté des Sciences Université Fédérale du Cameroun, 7-8: 97–140.
Dragesco, J., and Dragesco-Kernéis, A., 1986. Ciliés libres de l’Afrique intertropicale. Introduction à la connaissance et à l’étude des ciliés. Faune tropicale (Éditions de l’OSTROm), 26: 1–559.
Fenchel, T., and Finlay, B. J., 1984. Geotaxis in the ciliated protozoon Loxodes. Journal of Experimental Biology, 110 (1): 17–33.
Fenchel, T., and Finlay, B. J., 1986. The structure and function of Müller vesicles in loxodid ciliates. The Journal of Protozoology, 33 (1): 69–76.
Finlay, B. J., 1985. Nitrate respiration by protozoa (Loxodes spp.) in the hypolimnetic nitrite maximum of a productive freshwater pond. Freshwater Biology, 15 (3): 333–346.
Finlay, B. J., Fenchel, T., and Gardener, S., 1986. Oxygen perception and O2 toxicity in the freshwater ciliated protozoon Loxodes. Journal of Eukaryotic Microbiology, 33 (2): 157–165.
Foissner, W., and Rieder, N., 1983. Light and scanning electron-microscopical studies about the infraciliature of Loxodes striatus (Engelmann, 1862) and Loxodes magnus Stokes, 1887 (Protozoa, Ciliophora). Zoologischer Anzeiger, 210 (1-2): 3–13.
Foissner, W., 1995. Tropical protozoan diversity: 80 ciliate species (Protozoa, Ciliophora) in a soil sample from a tropical dry forest of Costa Rica, with descriptions of four new genera and seven new species. Archiv für Protistenkunde, 145 (1-2): 37–79.
Foissner, W., Berger, H., Blatterer, H., and Kohmann, F., 1995. Taxonomische und ökologische Revision der Ciliaten des Saprobiensystems. In: Band IV: Gymnostomatea, Loxodes, Suctoria. Bayerisches Landesamt für Wasserwirtschaft, München, 502pp.
Foissner, W., 1996. A redescription of Remanella multinucleata (Kahl, 1933) nov. gen., nov. comb. (Ciliophora, Karyorelictea), emphasizing the infraciliature and extrusomes. European Journal of Protistology, 32 (2): 234–250.
Foissner, W., and Al-Rasheid, K. A. S., 1999. Updating the trachelocercids (Ciliophora, Karyorelictea). VI. A detailed description of Sultanophrys arabica nov. gen., nov. spec. (Sultanophryideae nov. fam.). European Journal of Protistology, 35: 146–160.
Gao, F., Katz, L. A., and Song, W., 2013. Multigene-based analyses on evolutionary phylogeny of two controversial ciliate orders: Pleuronematida and Loxocephalida (Protista, Ciliophora, Oligohymenophorea). Molecular Phylogenetics & Evolution, 68 (1): 55–63.
Gao, F., Warren, A., Zhang, Q., Gong, J., Miao, M., Sun, P., Xu, D., Huang, J., Yi, Z., and Song, W., 2016. The all-data-based evolutionary hypothesis of ciliated protists with a revised classification of the phylum Ciliophora (Eukaryota, Alveolata). Scientific Reports, 6: 24874, DOI: 10.1038/srep24874.
Goulder, R., 1972. Grazing by the ciliated protozoon Loxodes magnus on the alga Scenedesmus in a eutrophic pond. Oikos, 23 (1): 109–115.
Hall, T. A., 1999. BioEdit: A user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucleic Acids Symposium Series, 41: 95–98.
Hines, H. N., Mccarthy, P. J., and Esteban, G. F., 2016. The first record for the Americas of Loxodes rex, a flagship ciliate with an alleged restricted biogeography. Microbial Ecology, 71 (1): 5–8.
Kim, J. H., Kwon, C. B., Yoon, J. S., and Shin, M. K., 2009. First record of three Loxodes ciliates (Ciliophora: Loxodida: Loxodidae) from Korea. Korean Journal of Systematic Zoology, 25 (3): 301–308.
Liu, W. W., Jiang, J. M., Xu, Y., Pan, X. M., Qu, Z. S., Luo, X. T., Warren, A., Ma, H. G., and Pan, H. B., 2017. Great diversity in marine ciliates: Fauna studies in China seas during the years 2011–2016. European Journal of Protistology, 61: 424–438.
Lynn, D. H., 2008. The Ciliated Protozoa: Characterization, Classification, and Guide to the Literature. 3rd edition, Springer, Dordrecht, 628pp.
Medlin, L., Elwood, H. J., Stickel, S., and Sogin, M. L., 1988. The characterization of enzymatically amplified eukaryotic 16S-like rDNA-coding regions. Gene, 71 (2): 491–499.
Miller, M. A., Pfeiffer, W., and Schwartz, T., 2010. Creating the CIPRES Science Gateway for inference of large phylogenetic trees. Gateway Computing Environments Workshop. New Orleans, LA, 1–8, DOI: 10.1109/GCE.2010.5676129.
Müller, O. F., 1786. Animalcula Infusoria Fluviatilia et Marina, quae Detexit, Systematice Descripsit et ad Vivum Delineari Curavit. Mölleri, Hauniae, 367pp.
Neugebauer, D. C., Machemer-Röhnisch, S., Nagel, U., Bräucker, R., and Machemer, H., 1998. Evidence of central and peripheral gravireception in the ciliate Loxodes striatus. Journal of Comparative Physiology A, 183 (3): 303–311.
Nylander, J. A. A., 2004. MrModeltest version 2. Evolutionary Biology Center, Uppsala University, Uppsala.
Raikov, I. B., 1959. Der Formwechsel des Kernapparate einiger niederer Ciliaten. II. Die Gattung Loxodes. Archiv für Protistenkunde, 104: 1–42.
Raikov, I. B., 1982. The Protozoan Nucleus–Morphology and Evolution. Springer-Verlag, 474pp.
Raikov, I. B., 1994. The nuclear apparatus of some primitive ciliates, the karyorelictids: Structure and divisional reorganization. Italian Journal of Zoology, 61 (1): 19–28.
Raikov, I. B., 1996. Nuclei of ciliates. In: Ciliates: Cells as Organisms. Hausmann, K., and Bradbury, P. C., eds., Fischer Verlag, Stuttgart, 221–242.
Sela, I., Ashkenazy, H., Katoh, K., and Pupko, T., 2015. GUIDANCE2: Accurate detection of unreliable alignment regions accounting for the uncertainty of multiple parameters. Nucleic Acids Research, 43 (W1): W7-W14.
Song, W., Warren, A., and Hu, X., 2009. Free-Living Ciliates in the Bohai and Yellow Seas, China. Science Press, Beijing, 520pp.
Stamatakis, A., Hoover, P., and Rougemont, J., 2008. A rapid bootstrap algorithm for the RAxML web servers. Systematics and Biodiversity, 57: 758–771.
Wang, C., Zhang, T., Wang, Y., Katz, L. A., Gao, F., and Song, W., 2017. Disentangling sources of variation in SSU rDNA sequences from single cell analyses of ciliates: Impact of copy number variation and experimental error. Proceedings Biological Sciences, 284 (1859): 20170425.
Wilbert, N., 1975. Eine verbesserte Technik der Protargolimprägnation für Ciliaten. Mikrokosmos, 64: 171–179.
Xu, Y., Fan, X. P., Al-Farraj, S. A., and Hu, X., 2017. Morphological description of two new marine ciliates (Ciliophora, Karyorelictea, Cryptopharyngidae): Apocryptopharynx discoidalis spec. nov. and Cryptopharynx minutus spec. nov. European Journal of Protistology, 58: 77–86.
Xu, Y., Gao, S., Hu, X., Al-Rasheid, K. A., and Song, W., 2013a. Phylogeny and systematic revision of the karyorelictid genus Remanella (Ciliophora, Karyorelictea) with descriptions of two new species. European Journal of Protistology, 49 (3): 438–452.
Xu, Y., Li, J. M., Song, W., and Warren, A., 2013b. Phylogeny and establishment of a new ciliate family, Wilbertomorphidae fam. nov. (Ciliophora, Karyorelictea), a highly specialized taxon represented by Wilbertomorpha colpoda gen. nov., spec. nov. Journal of Eukaryotic Microbiology, 60: 480–489.
Xu, Y., Miao, M., Warren, A., and Song, W., 2012. Diversity of the karyorelictid ciliates: Remanella (Protozoa, Ciliophora, Karyorelictida) inhabiting intertidal areas of Qingdao, China, with morphological and molecular information on three species. Systematics & Biodiversity, 10: 207–219.
Xu, Y., Pan, H., Miao, M., Hu, X., Al-Farraj, S. A., Al-Rasheid, K. A., and Song, W., 2015. Morphology and phylogeny of two species of Loxodes (Ciliophora, Karyorelictea), with description of a new subspecies, Loxodes striatus orientalis subsp. n. Journal of Eukaryotic Microbiology, 62 (2): 206–216.
Xu, Y., Shao, C., Miao, M., and Song, W., 2013c. Redescription of Parasonderia vestita (Kahl, 1928) comb. nov. (Ciliophora, Plagiopylida), with notes on its phylogeny based on SSU rRNA gene. European Journal of Protistology, 49: 106–113.
Xu, Y., Yan, Y., Li, L., Al-Rasheid, K. A. S., Al-Farraj, S. A., and Song, W., 2014. Morphology and phylogeny of three karyorelictean ciliates (Protista, Ciliophora), including two novel species, Trachelocerca chinensis sp. n. and Tracheloraphis dragescoi sp. n. International Journal of Systematic & Evolutionary Microbiology, 64: 4084–4097.
Yan, Y., Chen, X. M., Chen, X. R., Gao, F., Al-Farraj, S. A., and Al-Rasheid, K. A. S., 2015a. Morphology and molecular phylogeny of three marine Condylostoma species from China, including two new ones (Ciliophora, Heterotrichea). European Journal of Protistology, 51: 66–78.
Yan, Y., Fan, Y., Chen, X. R., Li, L. F., Warren, A., Al-Farraj, S. A., and Song, W., 2016a. Taxonomy and phylogeny of three heterotrich ciliates (Protozoa, Ciliophora), with description of a new Blepharisma species. Zoological Journal of the Linnean Society, 177: 320–334.
Yan, Y., Gao, F., Xu, Y., Al-Rasheid, K. A. S., and Song, W., 2015b. Morphology and phylogeny of three trachelocercid ciliates, with description of a new species, Trachelocerca orientalis spec. nov. (Ciliophora, Karyorelictea). Journal of Eukaryotic Microbiology, 62: 157–166.
Yan, Y., Rogers, A. J., Gao, F., and Katz, L. A., 2017. Unusual features of non-dividing somatic macronuclei in the ciliate class Karyorelictea. European Journal of Protistology, 61: 399–408.
Yan, Y., Xu, Y., Al-Farraj, S. A., Al-Rasheid, K. A. S., and Song, W., 2016b. Morphology and phylogeny of three trachelocercids (Protozoa, Ciliophora, Karyorelictea), with description of two new species and insight to the evolution of the family Trachelocercidae. Zoological Journal of the Linnean Society, 177: 306–319.
Yan, Y., Xu, Y., Yi, Z., and Warren, A., 2013. Redescriptions of three trachelocercid ciliates (Protista, Ciliophora, Karyorelictea), with notes on their phylogeny based on small subunit rDNA gene sequences. International Journal of Systematic & Evolutionary Microbiology, 63: 3506–3514.