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The Resurrection of the Genus Bergia (Anthozoa, Zoantharia, Parazoanthidae)

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The Resurrection of the Genus Bergia (Anthozoa, Zoantharia, Parazoanthidae) Systematics and Biodiversity ISSN: 1477-2000 (Print) 1478-0933 (Online) Journal homepage: http://www.tandfonline.com/loi/tsab20 The resurrection of the genus Bergia (Anthozoa, Zoantharia, Parazoanthidae) Javier Montenegro, Martyn E. Y. Low & James Davis Reimer To cite this article: Javier Montenegro, Martyn E. Y. Low & James Davis Reimer (2015): The resurrection of the genus Bergia (Anthozoa, Zoantharia, Parazoanthidae), Systematics and Biodiversity, DOI: 10.1080/14772000.2015.1101028 To link to this article: http://dx.doi.org/10.1080/14772000.2015.1101028 View supplementary material Published online: 21 Dec 2015. Submit your article to this journal View related articles View Crossmark data Full Terms & Conditions of access and use can be found at http://www.tandfonline.com/action/journalInformation?journalCode=tsab20 Download by: [Ryukyu University] Date: 21 December 2015, At: 17:23 Systematics and Biodiversity (2015), 1À11 Research Article The resurrection of the genus Bergia (Anthozoa, Zoantharia, Parazoanthidae) JAVIER MONTENEGRO1,3, MARTYN E. Y. LOW2 & JAMES DAVIS REIMER1,3 1Molecular Invertebrate Systematics and Ecology Laboratory, Graduate School of Engineering and Science, University of the Ryukyus, Senbaru 1, Nishihara, Okinawa 903-0213, Japan 2Lee Kong Chian Natural History Museum, Faculty of Science, National University of Singapore, 2 Conservatory Drive, Singapore 117377, Republic of Singapore 3Tropical Biosphere Research Center, University of the Ryukyus, Senbaru 1, Nishihara, Okinawa 903-0213, Japan (Received 5 May 2015; accepted 20 August 2015) The genus Bergia was established by Duchassaing de Fonbressin and Michelotti in 1860, for two species, Bergia catelunaris and B. vialactea. Subsequently, in 1903 Duerden recognized these two species as conspecific, and used B. catelunaris in favour of B. vialactea, and transferred B. catelunaris to the genus Parazoanthus. However, over the last decade, it has been found that the genus Parazoanthus is actually polyphyletic and therefore it has gradually been divided and redefined. Based on phylogenetic analyses, Parazoanthus sensu stricto was recently limited to species which form associations with sponges, but it still comprised of three distinctive and monophyletic subclades. Of these clades, one Parazoanthus clade contains the type species for Parazoanthus, P. axinellae, while another clade was recently described as the genus Umimayanthus based on mitochondrial 16S-rDNA sequences. However, the other remaining Parazoanthus clade contains P. catenularis, the original type species of the genus Bergia. Based on a concatenated set of published data for five molecular markers (18S-rDNA, 28S-rDNA, ITS1/5.8S/ITS2-rDNA, 16S-rDNA and COI-DNA), this study confirms the monophyly of Bergia in the context of Parazoanthidae. The phylogenetic analyses strongly support the resurrection of the genus Bergia as a valid taxonomic unit. Key words: Bergia, Parazoanthidae, Parazoanthus, sponge association, Umimayanthus Introduction on the same sponge two fairly distinct types of growth could be present, and he therefore considered these two The genus Bergia was established by Duchassaing de species as conspecific. Duerden (1903) synonymized both Fonbressin and Michelotti (1860) and defined by having names, using Bergia catenularis in favour of Bergia via- “very short polyps forming a chain-like incrustation on Downloaded by [Ryukyu University] at 17:23 21 December 2015 lactea (Low & Reimer, 2011). Additionally, Duerden the surface of sponges, with polyps arising from one (1903) synonymized Bergia with Parazoanthus Haddon another by stolons (propagules), not from a common and Shackleton, 1891 (family Parazoanthidae Delage & membrane, and originating from the upper or cephalic Herouard, 1901) after concluding that Bergia catelunaris part of the polyp” (p. 54 translated by Duerden, 1903) and possessed all the important anatomical diagnostic charac- by having “commensalistic and cateniform habits of colo- teristics that defined the genus Parazoanthus. nization” (p. 54 translated by Duerden, 1903). More recently, based on phylogenetic analyses, the Bergia was originally established to contain two spe- genus Parazoanthus has undergone further modifications. cies, Bergia catelunaris and Bergia vialactea, but after Sinniger, Montoya-Burgos, Chevaldonne and Pawlowski inspecting a large number of specimens, Duerden (1903) (2005) used mitochondrial DNA sequences and found that concluded that “it was very doubtful that any specific sep- the genus Parazoanthus was a polyphyletic group consist- aration can be maintained given the great variability on ing of several independent and well-supported subclades. habits according to age” (p. 26) and based on the fact that In following publications the genus was subsequently revised; Parazoanthus sensu stricto was defined as para- Correspondence to: Javier Montenegro. E-mail: jmontzalez@ zoanthids living on sponges, the genus Antipathozoanthus gmail.com Sinniger, Reimer & Pawlowski, 2010 was erected within ISSN 1477-2000 print / 1478-0933 online Ó The Trustees of the Natural History Museum, London 2015. All Rights Reserved. http://dx.doi.org/10.1080/14772000.2015.1101028 2 J. Montenegro et al. Parazoanthidae for zoantharian species associated with re-established as a valid genus within the family antipatharians, and the genus Hydrozoanthus Sinniger, Parazoanthidae. Reimer & Pawlowski, 2010 was erected within the family Hydrozoanthidae for species associated with hydrozoans. Nevertheless, under this new definition Parazoanthus was Materials and methods still polyphyletic and consisted of three distinctive mono- phyletic clades, designated A, B and C in Sinniger et al. Sequences (2010). Subsequently, Montenegro, Sinniger and Reimer Previously published sequences of family Parazoanthidae, (2015) used sequences from the nuclear partial ribosomal including the genera Isozoanthus, Savalia, Bullagummi- internal transcribed spacer 1, 5.8S rDNA, and partial zoanthus, Mesozoanthus, Kauluzoanthus, Hurlizoanthus, internal transcribed spacer 2 (ITS1/5.8S/ITS2-rDNA), Zibrowius, Corallizoanthus, Kulamanamana, Antipatho- nuclear asparagine-linked glycosylation 11 (ALG11- zoanthus, Bergia, Umimayanthus and Parazoanthus, and DNA), mitochondrial 16S ribosomal (16S-rDNA), and sequences of family Epizoanthidae, genus Epizoanthus, mitochondrial cytochrome c oxidase subunit 1 (COI- Gray, 1867, were downloaded from GenBank for 18S- DNA), combined with morphological characteristics, and rDNA, 28S-rDNA, ITS1/5.8S/ITS2-rDNA, 16S-rDNA split clade B from Parazoanthus into the genus Umi- and COI-DNA. For each marker shorter or poor quality mayanthus (see Montenegro et al., 2015). Thus, currently sequences were eliminated and a total of 198 sequences the genus Parazoanthus consists of two clades, with the were used. type species of Parazoanthus, P. axinellae (Schmidt, For the genus Epizoanthus 22 sequences were used; six 1862), within clade A, and P. catenularis (Duchassaing sequences for 18S-rDNA, three sequences for 28S-rDNA, de Fonbressing & Michelotti, 1860), the type species of three sequences for ITS1/5.8S/ITS2-rDNA, seven sequen- the genus Bergia Duchassaing de Fonbressin &Michelotti, ces for 16S-rDNA and three sequences for COI-DNA. 1860, included inside clade C. Inside family Parazoanthidae, for genus Bergia 23 Within the family Parazoanthidae 12 genera are cur- sequences were used: one for 18S-rDNA, three for 28S- rently recognized as valid (Table 1). In this study we use rDNA, nine for ITS1/5.8S/ITS2-rDNA, six for 16S- previously published and publicly available sequence data rDNA, and four for COI-DNA; for Umimayanthus 54 for all genera from five phylogenetic markers (18S-rDNA, sequences were used: two for 18S-rDNA, two for 28S- 28S-rDNA, ITS1/5.8S/ITS2-rDNA, 16S-rDNA and COI- rDNA, 19 for ITS1/5.8S/ITS2-rDNA, 15 for 16S-rDNA, DNA) to demonstrate that the genus Bergia is a well-sup- and 16 for COI-DNA; for Parazoanthus 53 sequences ported monophyletic taxon, separate from Parazoanthus were used: four for 18S-rDNA, eight for 28S-rDNA, 14 and all other parazoanthid genera, and therefore should be for ITS1/5.8S/ITS2-rDNA, 19 for 16S-rDNA and eight Table 1. Present classification of family Parazoanthidae. Diagnostic character(s) (Daly et al., 2007; Sinniger et al., 2010, Classification 2013; Montenegro et al., 2015) Zoantharia Gray, 1832 Clonal and soft-bodied polyps with two rows of marginal tentacles Downloaded by [Ryukyu University] at 17:23 21 December 2015 Macrocnemina Haddon & Shackleton, 1891 Fifth pair of mesenteries complete Parazoanthidae Delage & Herouard, 1901 Macrocnemic zoantharians with an endodermal sphincter muscle Antipathozoanthus Sinniger, Reimer & Pawlowski, 2010 DNA, grows exclusively on antipatharians Bullagummizoanthus Sinniger, Ocana~ & Baco, 2013 DNA, insertion/deletion pattern in 16S-rDNA V5 region Corallizoanthus Reimer in Reimer, Nonaka, Sinniger & Iwase, 2008 DNA, substrate specificity Hurlizoanthus Sinniger, Ocana~ & Baco, 2013 DNA, insertion/deletion pattern in 16S-rDNA V5 region Isozoanthus Carlgren, in Chun, 1903 No ring sinus, polyps solitary or weak coenenchyme Kauluzoanthus Sinniger, Ocana~ & Baco, 2013 DNA, insertion/deletion pattern in 16S-rDNA V5 region Kulamanamana Sinniger, Ocana~ & Baco, 2013 DNA, insertion/deletion pattern in 16S-rDNA V5 region Mesozoanthus Sinniger & Haussermann, 2009 DNA, absence of biological association Parazoanthus Haddon & Shackleton, 1891 Mesogleal lacuna and canal forming a ring sinus, substrate specificity (sponges) Savalia
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