Higher level taxonomy and molecular phylogenetics of the Choanoflagellatea

Journal of Eukaryotic Microbiology

Volumn 58, Issue 5, 2011, Pages 452-462

  Nitsche, Frank ^a   Carr, Martin ^b   Arndt, Hartmut ^a   Leadbeater, Barry S C ^c  

^a UNIVERSITY OF COLOGNE   (Germany)

^b UNIVERSITY OF LEEDS   (United Kingdom)

^c UNIVERSITY OF BIRMINGHAM   (United Kingdom)

Author keywords

Acanthoecida; choanoflagellates; Craspedida; Holozoa; taxa misidentification

Indexed keywords

RIBOSOME DNA; SEA WATER;

FLAGELLATE; MARINE ECOSYSTEM; MOLECULAR ANALYSIS; PHYLOGENETICS; PHYLOGENY; PROTEIN; TAXONOMY;

ARTICLE; CLASSIFICATION; EUKARYOTE; GENETICS; ISOLATION AND PURIFICATION; MOLECULAR GENETICS; NUCLEOTIDE SEQUENCE; PARASITOLOGY; PHYLOGENY;

DNA, RIBOSOMAL; EUKARYOTA; MOLECULAR SEQUENCE DATA; PHYLOGENY; SEAWATER;

ACANTHOECIDAE; CHOANOFLAGELLIDA; CODONOSIGIDAE; DERRIS TRIFOLIATA; HOLOZOA (GROUP); LORICA; METAZOA; SALPINGOECIDAE;

EID: 80053178237     PISSN: 10665234     EISSN: 15507408     Source Type: Journal    
DOI: 10.1111/j.1550-7408.2011.00572.x     Document Type: Article

References (96)

1. Adl, S. M., Simpson, A. G. B., Farmer, M. A., Andersen, R. A., Anderson, O. R., Barta, J. R., Bowser, S. S., Brugerolle, G., Fensome, R. A., Fredericq, S., James, T. Y., Karpov, S., Kugrens, P., Krug, J., Lane, C. E., Lewis, L. A., Lodge, J., Lynn, D. H., Mann, D. G., McCourt, R. M., Mendoza, L., Moestrup, Ø., Mozley-Standridge, S. E., Nerad, T. A., Shearer, C. A., Smirnov, A. V., Spiegel, F. W., &, Taylor, M. F. J. R., 2005. The new higher level classification of eukaryotes with emphasis on the taxonomy of protists. J. Eukaryot. Microbiol., 52: 399-451. (Pubitemid 43923926)
2. Baker, G. C., Beebee, T. J. C., &, Ragan, M. A., 1999. Prototheca richardsi, a pathogen of anuran larvae, is related to a clade of protistan parasites near the animal-fungal divergence. Microbiology, 145: 1777-1784. (Pubitemid 29352022)
3. Benny, G. L., &, O'Donnell, K., 2000. Amoebidium parasiticum is a protozoan, not a trichomycete. Mycologia, 92: 1133-1137.
4. Boenigk, J., &, Arndt, H., 2000. Comparative studies on the feeding behavior of two heterotrophic nanoflagellates: the filter-feeding choanoflagellate Monosiga ovata and the raptorial-feeding kinetoplastid Rhynchomonas nasuta. Aquat. Microb. Ecol., 22: 243-249. (Pubitemid 34235208)
5. Bourrelly, P., 1968. Les Algues D'eau Douce. Initiation á La Systématique. Tome II: Les Algues Jaunes et Brunes. N. Boubée et Cie, Paris. 438p.
6. Buck, K. R., Marchant, H. J., Thomsen, H. A., &, Garrison, D. L., 1990. Kakoeca antarctica gen. et sp. n., a loricate choanoflagellate (Acanthoecidae, Choanoflagellida) from Antarctic sea ice with a unique protoplast suspensory membrane. Zool. Scr., 19: 389-394.
7. Carr, M., Leadbeater, B. S. C., &, Baldauf, S. L., 2010. Conserved meiotic genes point to sex in the choanoflagellates. J. Eukaryot. Microbiol., 57: 56-65.
8. Carr, M., Cotton, S., Földvári, M., &, Kotrba, M., 2006. A description of a new species of Diasemopsis (Diptera, Diopsidae) from the Comoro Islands with morphological, molecular and allometric data. Zootaxa, 1211: 1-19. (Pubitemid 43926420)
9. Carr, M., Nelson, M., Leadbeater, B. S. C., &, Baldauf, S. L., 2008a. Three families of LTR retrotransposons are present in the genome of the choanoflagellate Monosiga brevicollis. Protist, 159: 579-590.
10. Carr, M., Leadbeater, B. S. C., Hassan, R., Nelson, M., &, Baldauf, S. L., 2008b. Molecular phylogeny of choanoflagellates, the sister group to Metazoa. Proc. Natl. Acad. Sci. USA, 105: 16641-16646.
11. Cavalier-Smith, T., 1993. Kingdom Protozoa and its 18 phyla. Microbiol. Rev., 57: 953-994. (Pubitemid 24004511)
12. Cavalier-Smith, T., 1997. Amoeboflagellates and mitochondrial cristae in eukaryote evolution: megasystematics of the new protozoan subkingdoms Eozoa and Neozoa. Arch. Protistenkd., 147: 237-258.
13. Cavalier-Smith, T., 1998. Neomonada and the origin of animals and fungi. In:, Coombs, G. H., Vickerman, K., Sleigh, M. A., &, Warren, A., (ed.), Evolutionary Relationships Among Protozoa. Kluwer, London. p. 375-407.
14. Cavalier-Smith, T., 2009. Megaphylogeny, cell body plans, adaptive zones: causes and timing of eukaryote basal radiations. J. Eukaryot. Microbiol., 56: 26-33.
15. Cavalier-Smith, T., &, Chao, E. E. Y., 2003. Phylogeny of Choanozoa, Apusozoa, and other protozoa and early eukaryotic megaevolution. J. Mol. Evol., 56: 540-563. (Pubitemid 36418983)
16. Cavalier-Smith, T., Chao, E. E. Y., &, Oates, B., 2004. Molecular phylogeny of Amoebozoa and the evolutionary significance of the unikont Phalansterium. Eur. J. Protistol., 40: 21-48. (Pubitemid 38836714)
17. Chadefaud, M., 1960. Les végétaux non vasculaires (Cryptogamie). In:, Chadefaud, M., &, Emberger, L., (ed.), Traité de Botanique, Systématique. Masson et Cie, Paris. p. 1-1018.
18. Collins, A. G., 1998. Evaluating multiple alternative hypotheses for the origin of Bilateria: an analysis of 18S rRNA molecular evidence. Proc. Natl. Acad. Sci. USA, 95: 15458-15463. (Pubitemid 29018720)
19. Conejo, M., Bertin, M., Pomponi, S. A., &, Ellington, W. R., 2008. The early evolution of the phosphagen kinases-insights from choanoflagellates and poriferan arginine kinases. J. Mol. Evol., 66: 11-20.
20. Criscione, C. D., Watral, V., Whipps, C. M., Blouin, M. S., Jones, S. R. M., &, Kent, M. L., 2002. Ribosomal DNA sequences indicate isolated populations of Ichthyophonus hoferi in geographic sympatry in the north-eastern Pacific Ocean. J. Fish Dis., 25: 575-582. (Pubitemid 35301917)
21. Deflandre, G., 1960. Sur la presence de Parvicorbicula n. g. socialis (Meunier) dans le plancton de l'Antarctique (Terre Adelie). Revue Algologique New Ser, 5: 183-188.
22. Doweld, A., 2003. Diplosoeca-a new generic name in Craspedophyceae. Byulleten' Moskovskogo Obshchestva Ispytatelei Prirody Otdel Biologicheskii (Bull. Mosc. Soc. Natur. Biol. Ser.), 108: 59-60.
23. Edgar, R. C., 2004. MUSCLE: multiple sequence alignment with high accuracy and high throughput. Nucleic Acids Res., 32: 1792-1797. (Pubitemid 38832724)
24. Ehrenberg, C. G., 1831. Über die Entwickelung und Lebensdauer der Infusionthiere: nebst ferneren Beiträgen zu einer Vergleichung ihrer organischen Système. Abh. Königl. Akad. Wiss. Berlin, 1-154.
25. Ehrenberg, C. G., 1838. Die Infusionsthierchen als vollkommene Organismen. Voss, Leipzig.
26. Ellis, W. N., 1929. Recent researches on the Choanoflagellata (Craspedomonadines). Ann. Soc. R. Zool. Belg., 60: 49-88.
27. Espeland, G., &, Throndsen, J., 1986. Flagellates from Kils-fjorden, southern Norway, with description of two new species of Choanoflagellida. Sarsia, 71: 209-226.
28. Fairclough, S. R., Dayel, M. J., &, King, N., 2010. Multicellular development in a choanoflagellate. Curr. Biol., 20: R875-876.
29. Francé, R. H., 1897. A craspedomonadinák szervezete. Der organismus der Craspedomonaden.: Kir. M. Termeszéttudományi Társulat, Budapest. p. 1-248.
30. Frenzel, J., 1891. Über einige merkwürdige Protozoen Argentiniens. Z. Wiss. Zool., 53: 332-360.
31. Hara, S., &, Takahashi, E., 1984. Re-investigation of Polyoeca dichotoma and Acanthoeca spectabilis (Acanthoecidae: Choanoflagellida). J. Mar. Biol. Assoc. UK, 64: 819-827.
32. Hara, S., &, Takahashi, E., 1987. An investigation with the electron microscope of marine choanoflagellates (Protozoa: Choanoflagellida) from Osaka Bay, Japan. II. Two new genera and a new species of Acanthoecidae. Bull. Plankton. Soc. Jpn., 34: 15-23.
33. Hendy, M. D., &, Penny, D., 1989. A framework for the quantitative study of evolutionary trees. Syst. Zool., 38: 297-309.
34. Herr, R. A., Ajello, L., Taylor, J. W., Arseculeratne, S. N., &, Mendoza, L., 1999. Phylogenetic analysis of Rhinosporidium seeberi's 18S small-subunit ribosomal DNA groups this pathogen among members of the protoctistan Mesomycetozoa clade. J. Clin. Microbiol., 37: 2750-2754. (Pubitemid 29391881)
35. Hertel, L. A., Bayne, C. J., &, Loker, E. S., 2002. The symbiont Capsaspora owczarzaki, nov. gen. nov. sp., isolated from three strains of the pulmonate snail Biomphalaria glabrata is related to members of the Mesomycetozoea. Int. J. Parasitol., 32: 1183-1191. (Pubitemid 34761912)
36. Hibberd, D. J., 1976. The ultrastructure and taxonomy of the Chrysophyceae and Prymnesiophyceae (Haptophyceae): a survey with some new observations on the ultrastructure of the Chrysophyceae. Bot. J. Linn. Soc., 72: 55-80.
37. Hibberd, D. J., 1983. Ultrastructure of colonial colourless zooflagellates Phalansterium digitatum Stein (Phalansteriida ord. nov.) and Spongomonas uvella Stein (Spongomonadida ord. nov.). Protistologica, 19: 523-535.
38. Iyengar, M. O. P., &, Ramanathan, K. R., 1940. Cladospongia, a new member of the Craspedomonadaceae from Madras. J. Indian Bot. Soc., 19: 241-245.
39. James-Clark, H., 1866. Note on the Infusoria Flagellata and the Spongiae Ciliatae. Am. J. Sci., 1: 113-114.
40. James-Clark, H., 1867. On the Spongiae Ciliatae as Infusoria Flagellata; or observations on the structure, animality and relationship of Leucosolenia botryoides, Bowerbank. Ann. Mag. Nat. Hist., 1: 133-142.
41. Karpov, S. A., &, Leadbeater, B. S. C., 1998. Cytoskeleton structure and composition in choanoflagellates. J. Eukaryot. Microbiol., 45: 361-367. (Pubitemid 28281556)
42. Kent, W. S., 1878. A new field for the microscopist. Popular Sci. Rev., 2 (New Series): 113-132.
43. Kent, W. S., 1880-1882. In:, Bogue, D., (ed.), A Manual of the Infusoria. Vol. 1-3, London, 903 pp.
44. King, N., Westbrook, M. J., Young, S. L., Kuo, A., Abedin, M., Chapman, Fairclough, S., Hellsten, U., Isogai, Y., Letunic, I., Marr, M., Pincus, D., Putnam, D., Rokas, A., Wright, K. J., Zuzow, R., Dirks, W., Good, M., Goodstein, D., Lemons, D., Li, W., Lyons, J. B., Morris, A., Nichols, S., Richter, D. J., &, Salamov, A., JGI Sequencing, Bork, P., Lim, W. A., Manning, G., Miller, W. T., McGinnis, W., Shapiro, H., Tjian, R., Grigoriev, I. V., &, Rokhsar, D., 2008. The genome of the choanoflagellate Monosiga brevicollis and the origin of metazoans. Nature, 451: 783-788. (Pubitemid 351253174)
45. Klug, G., 1936. Salpingorhiza gen. nov. eine neue Crapsedomonadaceengattung. (Studien uber farblose Flagellaten II). Arch. Protistenkd., 87: 117-122.
46. Krylov, M. V., Dobrovoisky, A. A., Issi, I. V., Michalevich, V. I., Podlipaev, C. A., Reshetnyak, V. V., Seravin, L. I., Starobogatov, V. I., Shulman, C. C., &, Yankovsky, A. V., 1980. New advances in the system of lower organisms. Proc. Zool. Inst. Acad. Sci. USSR, 94: 122-132.
47. Lackey, J. B., 1940. Some new flagellates from the Woods Hole area. Am. Midl. Nat., 23: 463-471.
48. Lackey, J. B., 1942. Two new flagellate protozoa from the Tennessee River. Trans. Am. Microsc. Soc., 61: 36-39.
49. Lara, E., Moreira, D., &, López-García, P., 2010. The environmental clade LKM11 and Rozella form the deepest branching clade of fungi. Protist, 161: 116-121.
50. Lauterborn, R., 1894. Über die Winterfauna einiger Gewässer der Obertrheinebene mit Beschreibungen neuer Protozoën. Biologisches Centralblat, 14: 390-398.
51. Leadbeater, B. S. C., 1978. Renaming of Salpingoeca sensu Grøntved. J. Mar. Biol. Assoc. UK., 8: 511-515.
52. Leadbeater, B. S. C., 1980. Four new species of loricate choanoflagellates from South Brittany, France. Cah. Biol. Mar., 21: 345-353.
53. Leadbeater, B. S. C., 2008. Choanoflagellate lorica construction and assembly: the nudiform condition. I. Savillea species. Protist, 159: 259-268.
54. Leadbeater, B. S. C., 2010. Choanoflagellate lorica construction and assembly: the tectiform condition. Volkanus costatus (=Diplotheca costata). Protist, 161: 160-176.
55. Leadbeater, B. S. C., &, Cheng, R., 2010. Costal strip production and lorica assembly in the large tectiform choanoflagellate Diaphanoeca grandis Ellis. Eur. J. Protistol., 46: 96-110.
56. Leadbeater, B. S. C., Henouil, M., &, Berovic, N., 2008a. Choanoflagellate lorica construction and assembly: the nudiform condition. II. Acanthoeca spectabilis. Protist, 159: 495-505.
57. Leadbeater, B. S. C., Hassan, R., Nelson, M., Carr, M., &, Baldauf, S. L., 2008b. A new genus, Helgoeca gen. nov., for a nudiform choanoflagellate. Eur. J. Protistol., 44: 227-237.
58. Lee, J. J., Hutner, S. H., &, Bovee, E. C., 1985. An Illustrated Guide to the Protozoa. Society of Protozoologists. Vol. 1, Lawrence, KS, 689 pp.
59. Lepère, C., Domaizon, I., &, Debroas, D., 2008. Unexpected importance of potential parasites in the composition of the freshwater small-eukaryote community. Appl. Environ. Microbiol., 74: 2940-2949. (Pubitemid 351746995)
60. Loeblich III, A. R., 1967. Notes on the divisions Chlorophyta, Chrysophyta, Pyrrophyta and Xanthophyta and the family Parastigaceae. Taxon, 16: 230-236.
61. Logares, R., BrÃte, J., Bertilsson, S., Clasen, J. L., Shachian-Tabrizi, K., &, Rengefors, K., 2009. Infrequent marine-freshwater transitions in the microbial world. Trends Microbiol., 17: 414-422.
62. Manton, I., &, Bremer, G., 1981. Observations on lorica structure and aspects of replication in Pleurasiga sphyrelata Thomsen complex (=Polyfibula spp., gen. n.) (Choanoflagellata). Zool. Scripta, 10: 273-291.
63. Manton, I., Bremer, G., &, Oates, K., 1981. Problems of structure and biology in a large collared flagellate (Diaphanoeca grandis Ellis) from Arctic seas. P. Roy. Soc. Lond. B. Bio, 213: 15-26. (Pubitemid 12225200)
64. Marchant, H. J., &, Perrin, R., 1986. Planktonic choanoflagellates from two Antarctic lakes including a description of Spiraloecion didymocostatum gen. et sp. nov. Polar Biol., 5: 207-210.
65. Medina, M., Collins, A. G., Taylor, J. W., Valentine, J. W., Lipps, J. H., Amaral-Zettler, L., &, Sogin, M. L., 2003. Phylogeny of Opisthokonta and the evolution of multicellularity and complexity in Fungi and Metazoa. Int. J. Astro., 2: 203-211.
66. Medlin, L., Elwood, H. J., Stickel, S., &, Sogin, M. L., 1988. The characterization of enzymatically amplified eukaryotic 16S-like rRNA-coding regions. Gene, 71: 491-499. (Pubitemid 19010867)
67. Nedelcu, A. M., Blakney, A. J., &, Logue, K. D., 2009. Functional replacement of a primary metabolic pathway via independent eukaryote-to- eukaryote gene transfers and selective retention. J. Evol. Biol., 22: 1882-1894.
68. Nedelcu, A. M., Miles, I. H., Karol, K., &, Fagir, A., 2008. Adaptive eukaryote-to-eukaryote gene transfer: stress-related genes of algal origin in the closest unicellular relatives of animals. J. Evol. Biol., 21: 1852-1860.
69. Nitsche, F., &, Arndt, H., 2008. A new choanoflagellate species from Taiwan: morphological and molecular biological studies of Diplotheca elongata nov. spec. and D. costata. Eur. J. Protistol., 44: 220-226.
70. Norris, R. E., 1965. Neustonic marine Craspedomonadales (choanoflagellates) from Washington and California. J. Protozool., 12: 589-602.
71. Pereira, C. N., Di Rosa, I., Fagotti, A., Simoncelli, F., Pascolini, R., &, Mendozam, L., 2005. The pathogen of frogs Amphibiocystidium ranae is a member of the order Dermocystida in the class Mesomycetozoea. J. Clin. Microbiol., 43: 192-198. (Pubitemid 40096300)
72. Philippe, H., Snell, E. A., Bapteste, E., Lopez, P., Holland, P. W. H., &, Casane, D., 2004. Phylogenomics of eukaryotes: impact of missing data on large alignments. Mol. Biol. Evol., 21: 1740-1752. (Pubitemid 39096890)
73. Quintela-Alonso, P., Nitsche, F., &, Arndt, H., 2011. Molecular characterization and revised systematics of Microdiaphanosoma arcuatum (Ciliophora, Colpodea). J. Eukaryot. Microbiol., 58: 114-119.
74. Ragan, M. A., Goggins, C. L., Cawthorn, R. J., Cerenius, L., Jamienson, A. V. C., Plourde, S. M., Rand, T. G., Söderhäll, K., &, Gutell, R. R., 1998. A novel clade of protistan parasites near the animal-fungal divergence. Proc. Natl. Acad. Sci. USA, 93: 11907-11912. (Pubitemid 26347222)
75. Ronquist, F., &, Huelsenbeck, J. P., 2003. MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics B, 19: 1572-1574.
76. Ruiz-Trillo, I., Roger, A. J., Burger, G., Gray, M. W., &, Lang, B. F., 2008. A phylogenomic investigation into the origin of Metazoa. Mol. Biol. Evol., 25: 664-672. (Pubitemid 351398321)
77. Schiller, J., 1925. Die planktonischen Vegetationen des Adriatischen Meeres. B. Chrysomonadina, Heterokontae, Cryptomonadina, Eugleninae, Volvocales. I. Systematischer Teil. Arch. Protistenkd., 53: 59-123.
78. Schiller, J., 1953. Über neuen Craspedomonaden (Choanoflagellaten). Arch. Hydrobiol., 48: 248-59.
79. Senn, G., 1900. In:, Flagellaten. Engler, H. G. A., &, Prantl, K. A. E., (ed.), Die Natürlichen Pflanzenfamilien. Vol 1, Part 1a. Wilhelm Engelmann Verlag, Leipzig. p. 93-188.
80. Snell, E. A., Furlong, R. F., &, Holland, P. W. H., 2001. Hsp70 sequences indicate that choanoflagellate are closely related to animals. Curr. Biol., 11: 967-970. (Pubitemid 32589188)
81. Snell, E. A., Brooke, N. M., Taylor, W. R., Casane, D., Philippe, H., &, Holland, P. W. H., 2006. An unusual choanoflagellate protein released by Hedgehog autocatalytic processing. P. Roy. Soc. Lond. B. Bio., 273: 401-407. (Pubitemid 46780546)
82. Stamatakis, A., 2006. RAxML-VI-HPC: maximum likelihood-based phylogenetic analyses with thousands of taxa and mixed models. Bioinformatics, 22: 2688-2690. (Pubitemid 44642609)
83. Steenkamp, E. T., Wright, J., &, Baldauf, S. L., 2006. The protistan origins of animals and fungi. Mol. Biol. Evol., 23: 93-106. (Pubitemid 41785957)
84. Stein, F. R., 1878. Der Organismus der Infusionthiere. Vol. III. Abt. W. Englemann, Leipzig.
85. Suga, H., Sasaki, G., Kuma, K., Nishiyori, H., Hirose, N., Su, Z., Iwabe, N., &, Miyata, T., 2008. Ancient divergence of animal protein tyrosine kinase genes demonstrated by a gene family tree including choanoflagellate genes. FEBS Lett., 582: 815-818.
86. Thomsen, H. A., 1976. Studies on marine choanoflagellates. II. Fine structural observations on some silicified choanoflagellates from the Isefjord (Denmark), including the description of two new species. Norweg. J. Bot., 23: 33-51.
87. Thomsen, H. A., 1979. Electron microscopical observations on brackish water nannoplankton from the Tvärminne area, S.W. Coast of Finland. Acta Bot. Fennica, 110: 11-37.
88. Thomsen, H. A., 1982. Planktonic choanoflagellates from Disco Bugt, West Greenland, with a survey of the marine nanoplankton of the area. Meddelelser om Gronland (Bioscience), 8: 3-35.
89. Thomsen, H. A., 1988. An electron microscopical study of marine loricate choanoflagellates: Nannoeca minuta (Leadbeater) gen. et comb. n. and Stephanoeca cupula (Leadbeater) comb. nov. Zool. Scr., 17: 315-23.
90. Thomsen, H. A., &, Boonruang, P., 1983a. A microscopical study of marine collared flagellates (Choanoflagellida) from the Andaman Sea, S.W. Thailand; species of Stephanacantha gen. nov.and Platypleura gen. nov. Protistologica, 19: 193-214.
91. Thomsen, H. A., &, Boonruang, P., 1983b. Ultrastructural observations on marine choanoflagellates (Choanoflagellida, Acanthoecidae) from the coast of Thailand; species of Apheloecion gen. nov. J. Plankton Res., 5: 729-753.
92. Thomsen, H. A., &, Boonruang, P., 1984. A light and electron microscopical investigation of loricate choanoflagellates (Choanoflagellida, Acanthoecidae) from the Andaman Sea, S.W. Thailand and Denmark; species of Cosmoeca gen. n. Zool. Sc., 13: 165-181.
93. Thomsen, H. A., &, Moestrup, Ø., 1984. Electron microscopical investigations on two loricate choanoflagellates (Choanoflagellida) Calotheca alata gen. et. sp. nov. and Syndetophyllum pulchellum gen. et. comb. nov. from Indo-Pacific localities. P. Roy. Soc. London B Bio., 219: 41-52.
94. Thomsen, H. A., Østergaard, J. B., &, Hansen, L. E., 1995. Loricate choanoflagellates from West Greenland (August 1988) including the description of Spinoeca buckii gen. et sp. nov. Europ. J. Protistol., 31: 38-44.
95. Throndsen, J., 1974. Planktonic choanoflagellates from North Atlantic waters. Sarsia, 56: 95-122.
96. Wainright, P. O., Hinkle, G., Sogin, M. L., &, Stickel, S. K., 1993. Monophyletic origins of the Metazoa: an evolutionary link with fungi. Science, 260: 340-342. (Pubitemid 23160139)