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Gastropoda: Columbellidae) from Eastern Atlantic

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Gastropoda: Columbellidae) from Eastern Atlantic An assessment of the genus Columbella Lamarck, 1799 (Gastropoda: Columbellidae) from eastern Atlantic Valeria RUSSINI Giulia FASSIO Maria Vittoria MODICA Department of Biology and Biotechnologies “Charles Darwin”, Sapienza University of Rome, Viale dell’Università 32, I-00185 Roma (Italy) Marta J. deMAINTENON University of Hawaii at Hilo, 200 W. Kawili Street, Hilo, HI 96720 (United States) Marco OLIVERIO Department of Biology and Biotechnologies “Charles Darwin”, Sapienza University of Rome, Viale dell’Università 32, I-00185 Roma (Italy) [email protected] (corresponding author) Published on 30 June 2017 urn:lsid:zoobank.org:pub:57395522-BE12-40AC-B504-C448A31F8A8D Russini V., Fassio G., Modica M. V., deMaintenon M. J. & Oliverio M. 2017. — An assessment of the genus Colum- bella Lamarck, 1799 (Gastropoda: Columbellidae) from eastern Atlantic. Zoosystema 39 (2): 197-212. https://doi. org/10.5252/z2017n2a2 ABSTRACT Th ree species of the neogastropod genus Columbella Lamarck, 1799 are recognised from the northeastern Atlantic and the Mediterranean. One is the common Mediterranean C. rustica (Linnaeus, 1758), with paucispiral protoconch, extending its range in the Atlantic South to Senegal and North to Portugal. Columbella adansoni Menke, 1853, with multispiral protoconch is restricted to the Macaronesian ar- KEY WORDS chipelagoes. A third species, also with multispiral protoconch, from West Africa is recognised through Columbellidae, molecular methods, and the name C. xiphitella Duclos, 1840 is employed by correcting the original East Atlantic, erroneous locality (“Californie”) to Gabon. Except for protoconch features, no major morphological Mediterranean, lectotypifi cation, characters are available to separate the three species; however diagnostic species-level diff erences in DNA-Barcoding. specifi c positions in the cytochrome c oxidase I (COI) sequences are present between all three species. RÉSUMÉ Étude du genre Columbella Lamarck, 1799 (Gastropoda: Columbellidae) dans l’Est de l’océan Atlantique. Trois espèces du genre de néogastropode Columbella Lamarck, 1799 sont reconnues dans le nord est de l’Atlantique et en Méditerranée. L’une est courante en Méditerranée, C. rustica (Linnaeus, 1758), au protoconche paucispiralé : son aire de répartition s’étend en Atlantique du Sénégal au nord du Portugal. Columbella adansoni Menke, 1853, au protoconche multispiralé, se limite aux archipels Macaronésiens. Une troisième espèce, caractérisée également par un protoconche multispiralé, est ori- MOTS CLÉS ginaire d’Afrique de l’Ouest : elle est reconnue par des méthodes moléculaires ; le nom de C. xiphitella Columbellidae, Duclos, 1840 lui est attribué après correction de la localité originale erronée (« Californie ») en Gabon. Atlantique de l’est, Mis à part l’aspect du protoconche, aucun caractère morphologique majeur ne permet de séparer les Méditerranée, lectotypifi cation, trois espèces ; cependant des positions précises dans les séquences du cytochrome c oxidase I (COI) DNA-Barcoding. présentent des diff érences supportant des diagnoses spécifi ques. ZOOSYSTEMA • 2017 • 39 (2) © Publications scientifi ques du Muséum national d’Histoire naturelle, Paris. www.zoosystema.com 197 Russini V. et al. INTRODUCTION HCO2198-LCO1490, we employed HCO2198 with the primer mlCOIint-F (5’-GGWACWGGWTGAACWGT- Columbella Lamarck, 1799 s.s. (type species Voluta merca- WTAYCCYCC-3’) designed to amplify a shorter fragment toria Linnaeus, 1758) is a genus of columbellid neogastro- (c. 300 bp) and employed in metabarcoding works (Leray pods (dove shells) including 17 recognised species, mostly et al. 2013). PCR amplifi cations were performed with the from tropical America and the East Atlantic/Mediterranean following conditions: initial denaturation of 5’ at 94°C, (WoRMS: Bouchet & Gofas 2015). Based on Moolenbeek & 35 amplifi cation cycles (30”/94°C, 40”/48-52°C, 50”/72°C), Hoenselaar (1991), Oliverio (1995), Rolán (2005), and followed by a fi nal phase of 7’ at 72°C. PCR products were Rolán & Ryall (1999), two species are currently recorded in purifi ed by ExoSAP-IT protocol (USB Corporation, Ohio, the eastern Atlantic and the Mediterranean Sea: Columbella USA) and Sanger sequenced by Macrogen Inc. (Th e Neth- rustica (Linnaeus, 1758), ranging over the entire Mediter- erlands). Forward and reverse sequences were assembled, ranean Sea, and extending into the neighbouring Atlantic checked for contamination and edited with Geneious 4.8.5 southward to Senegal, and northward to Portugal (it is (Drummond et al. 2009). absent in Galicia); and Columbella adansoni Menke, 1853, described from Cape Verde islands, and assumed to occur SPECIES DELIMITATION IN COLUMBELLIDAE SWAINSON, 1840 across Macaronesia, from the Azores to the Canary Islands, A total of 106 COI sequences from columbellid specimens and along the West African coasts from Ghana to Angola ascribed to the genera Alia H. Adams & A. Adams, 1853, (Oliverio 1995; Rolán & Ryall 1999; Rolán 2005). Colum- Amphissa H. Adams & A. Adams, 1853, Euplica Dall, 1889, bella rustica has a paucispiral protoconch, indicating non- Graphicomassa Iredale, 1929, Indomitrella Oostingh, 1940, planktotrophic development (leci thotrophic, possibly entirely Mitrella Risso, 1826, Pyrene Röding, 1798, Sulcomitrella or mostly intracapsular), whereas Columbella adansoni has Kuroda, Habe & Oyama, 1971 and Zafra A. Adams, 1860 a multispiral protoconch, indicating planktotrophic larval (plus some labelled as “columbellid indet.”) were either development. Th is is the only consistent morphological diag- provided by Nicolas Puillandre (ID MNHN-IM) or were nostic feature for the two species, which are otherwise quite retrieved from the GenBank (see Table 4). Sequences from variable in shell sculpture, colour and pattern. Preliminary Cancellopollia sp. (Gastropoda, Buccinoidea, Buccinidae) to a study of the bearing of diff erent larval developmental (EU015666.1; voucher MNHN-IM-2009-17854), and Pisania strategies on the genetic structure of populations (Modica striata Duclos, 1840 (MNHN-IM-2009-30664, Gastropoda, et al. 2017), we decided to assay samples of Columbella Buccinoidea, Buccinidae) were retrieved from Genbank to be from the eastern Atlantic and the Mediterranean to test the used as outgroups. COI sequences were manually aligned and currently accepted species boundaries by molecular data. checked for stop codons; 16S sequences were aligned using Th erefore, we examined specimens collected from locali- MAFFT 7 (Katoh et al. 2002), using the Q-INS-i algorithm ties spanning as much as possible the known range for the (Katoh & Toh 2008), which accounts for secondary struc- genus in the eastern Atlantic. As a result, a third species of tures. Highly variable regions, resulting in gap-rich fragments Columbella was discovered. with ambiguous alignment, were discarded using Gblocks 0.91b (Castresana 2000). All alignments are available from the authors on request. MATERIAL AND METHODS To defi ne species, we used Automatic Barcode Gap Dis- covery (ABGD, available at http://wwwabi.snv.jussieu.fr/ Sampling locality data (Fig. 1), Identifi cation (ID) catalogue public/abgd/), a distance-based method designed to detect numbers of the vouchers, and GenBank accession numbers the so-called “barcode gap” in the distribution of pairwise are reported in Table 1. A total of 29 specimens from the distances estimated in a COI alignment (Puillandre et al. East Atlantic and the Mediterranean were assayed. Specimens 2012a, b), and the criteria of divergence and reciprocal mono- were sampled by SCUBA or snorkelling, and fi xed in 95 phyly (Knowlton 2000; Wheeler & Meier 2000; Reid et al. to 100% ethanol. Vouchers are stored in the malacological 2006; Malaquias & Reid 2009). Th e COI sequence align- collection at Department of Biology and Biotechnologies ments were processed in ABGD (excluding the outgroups) “Charles Darwin” (“La Sapienza” University of Rome) un- using the Kimura-2-parameter (K2p) model and the follow- der BAU ID numbers and at Muséum national d’Histoire ing settings: a prior for the maximum value of intraspecifi c naturelle (Paris) under MNHN ID numbers. Genomic divergence between 0.001 and 0.1, 25 recursive steps within DNA was extracted using a proteinase K-phenol-chloroform the primary partitions defi ned by the fi rst estimated gap, and protocol (Oliverio & Mariottini 2001). Th e DNA-barcode a gap width of 0.1. fragment of the mitochondrial cytochrome c oxidase I (COI) We ran ABGD on the whole columbellid dataset of 136 COI and part of the 16S rRNA were amplifi ed by PCR using the sequences, to defi ne partition scheme(s) based on distance universal primers LCO1490 and HCO2198 (Folmer et al. distribution. Th en, species hypotheses as derived from ABGD 1994) and 16SA (Palumbi et al. 2002) and CGLeuUUR were tested against taxonomic recognition for the assayed (Hayashi 2003), respectively. For some crucial specimens specimens and for phylogenetic congruence. Phylogenetic from West Africa, fi xed in alcohol but thereafter preserved analyses of the COI, 16S and combined sequence align- dried, which were unsuccessfully assayed with the pair ments were conducted using Maximum likelihood (ML: 198 ZOOSYSTEMA • 2017 • 39 (2) East Atlantic Columbella FRANCE ITALY 375 40° PORTUGAL GREECE Mediterranean Sea MAROCCO TUNISIA 250 LIBYA EGYPT SENEGAL 125 10°N GHANA CAMEROON 0° GABON 0 0.03 0.06 0.09 0.12 0.15 0.18 0.21 0.24 0.27 0.30 0.33 0.36 0.39 Atlantic Ocean 10°S FIG. 2. — Histogram of the distribution of the pairwise estimated
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