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Molecular Evidence for the Coexistence of Two Sibling Species in Pylaiella Littoralis (Ectocarpales, Phaeophyceae) Along the Brittany Coast

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Molecular Evidence for the Coexistence of Two Sibling Species in Pylaiella Littoralis (Ectocarpales, Phaeophyceae) Along the Brittany Coast See discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/295605392 Molecular evidence for the coexistence of two sibling species in Pylaiella littoralis (Ectocarpales, Phaeophyceae) along the Brittany coast Article in Journal of Phycology · January 2015 CITATIONS READS 0 23 5 authors, including: stéphane Mauger Aurélien De Jode Station Biologique de Roscoff Institut Méditerranéen de Biodiversité et d'… 41 PUBLICATIONS 1,450 CITATIONS 8 PUBLICATIONS 8 CITATIONS SEE PROFILE SEE PROFILE Line Le Gall Christophe Destombe Muséum National d'Histoire Naturelle Pierre and Marie Curie University - Paris 6 102 PUBLICATIONS 1,079 CITATIONS 85 PUBLICATIONS 1,650 CITATIONS SEE PROFILE SEE PROFILE Some of the authors of this publication are also working on these related projects: JOURNÉES DU GRAAL : Graphes et fouille de données pour la recherche environnementale : des données, des questions de recherche et de nouvelles hypotheses View project All in-text references underlined in blue are linked to publications on ResearchGate, Available from: Christophe Destombe letting you access and read them immediately. Retrieved on: 24 October 2016 J. Phycol. 51, 480–489 (2015) © 2015 Phycological Society of America DOI: 10.1111/jpy.12291 MOLECULAR EVIDENCE FOR THE COEXISTENCE OF TWO SIBLING SPECIES IN PYLAIELLA LITTORALIS (ECTOCARPALES, PHAEOPHYCEAE) ALONG THE BRITTANY COAST1 Alexandre Geoffroy, Stephane Mauger, Aurelien De Jode Sorbonne Universites, UPMC Universite Paris 06, UMI 3614 Evolutionary Biology and Ecology of Algae, Station Biologique de Roscoff, CS 90074, Roscoff 29688, France CNRS, UMI 3614 Evolutionary Biology and Ecology of Algae, Station Biologique de Roscoff, Roscoff 29688, France Line Le Gall Museum National d’Histoire Naturelle, Institut de Systematique, Evolution, Biodiversite, UMR 7205 CNRS-EPHE-MNHN-UPMC, Equipe Exploration, Especes, Evolution, case postale No 39, 57 rue Cuvier, 75231 Cedex 05 Paris, France and Christophe Destombe2 Sorbonne Universites, UPMC Universite Paris 06, UMI 3614 Evolutionary Biology and Ecology of Algae, Station Biologique de Roscoff, CS 90074, Roscoff 29688, France CNRS, UMI 3614 Evolutionary Biology and Ecology of Algae, Station Biologique de Roscoff, Roscoff 29688, France The great phenotypic variability and the lack of tide; PCA, principal component analysis; rbcL, ribu- diagnostic characters in the genus Pylaiella render lose-1,5-biphosphate carboxylase oxygenase large the systematic study of this genus problematic. In subunits this study, we investigated the diversity of Pylaiella littoralis along the Brittany (France) coast using a DNA barcoding multilocus approach with Most species descriptions in brown algal groups fol- mitochondrial (cox1, nad1, and atp9) and low the morphological species concept and the chloroplastic (rbcL and atpB) markers associated delimitation of species is therefore essentially based with a population genetics approach using 10 on discontinuities in morphological characters (May- microsatellite markers. In addition, spatio-temporal den 1997). However, because of phenotypic plasticity sampling was conducted along the Brittany coast. and the lack of diagnostic characters in some groups, We sampled 140 individuals from four sites located species delimitation using conventional methods is between Saint-Malo and Concarneau (380 km) from often problematic and has led to erroneous taxo- April to October. Mitochondrial sequence data nomic assignments (Tellier et al. 2009, Tronholm revealed the occurrence of two sibling species, with et al. 2010). Analyses of molecular data have become a minimum of 2.4% divergence between them. the closest proxy of species delimitation (for review, Microsatellite genotypic data congruently revealed Leliaert et al. 2014). The combination of phyloge- two well-supported clusters matching the two netic and population genetics approaches can help mitochondrial clades of Pylaiella. Although gene determine algal diversity and study boundaries flow is limited between species, occurrence of between closely related species. As a phylogenetic genetic admixtures in some populations suggested approach, DNA barcoding—based on discontinuity that reproductive isolation is not complete. Our in sequence variation as diagnostic character—has study highlighted the complementarity of barcoding proven to be an effective tool for delineating species and population genetics approaches to shed light on (Sites and Marshall 2003) and has been successfully the evolutionary processes that lead to speciation. applied in algae to aggregate individuals into differ- ent closely related species (Leliaert et al. 2014). The Key index words: gene flow; microsatellite; multilocus Pylaiella littoralis population genetic approach has been used in brown barcode; ; sibling species. algae to detect gene flow discontinuity among taxa Abbreviations: atp9, ATP synthase subunit 9; atpB, (Tellier et al. 2011) and to identify hybridization or ATPase beta subunit; cox1, cytochrome c oxidase introgression events (Engel et al. 2005). Population subunit I; FCA, factorial correspondence analysis; genetic data acquired from microsatellite loci are nad1, NADH dehydrogenase subunit 1; nt, nucleo- extremely useful to estimate the extent of gene flow between closely related species. 1Received 27 June 2014. Accepted 19 January 2015. The brown algal genus Pylaiella includes species 2 Author for correspondence: e-mail christophe.destombe@sb-ros characterized by a filamentous structure with various coff.fr. Editorial Responsibility: O. De Clerck (Associate Editor) branching patterns. As in most algal groups, system- 480 SIBLING SPECIES IN PYLAIELLA LITTORALIS 481 atics in Pylaiella has been centered on morphologi- two sibling species in Brittany under the name of cal species concept. However, Pylaiella species are P. littoralis. known for their disconcerting phenotypic plasticity and their lack of diagnostic characters, making spe- MATERIALS AND METHODS cies delimitation very problematic (Fritsch 1952, Bolton 1979a). More than 90 specific and infraspe- Sampling. In total, 140 specimens of P. littoralis were sam- cific names have been reported for this genus, of pled at four different localities (16–62 specimens per site) which only seven are currently accepted taxonomi- along the coast of Brittany, France (Fig. 1), from Saint Malo to Concarneau (~380 km coastline). Sampling was conducted cally (Guiry and Guiry 2014). Pylaiella littoralis from April to October (Table 1) to compare spring and (Linnaeus) Kjellman is the most common accepted autumn samples. In the laboratory, a fragment of tissue from species and has been reported worldwide in the lit- each individual was preserved in silica gel for molecular toral zones of cold and cold temperate coasts of analysis. In addition, at least one specimen from each site was both hemispheres (Muller€ and Stache 1989, Silva pressed and mounted on an herbarium sheet to preserve ana- et al. 1999). More than 50 forms or varieties have tomical features (vouchers were deposited in the herbarium of the Museum National d’Histoire Naturelle, Paris, France, nos. been described on the basis of morphology, habitat, PC0533607-PC0533613). and seasonal distribution characteristics (Guiry and To complete the sampling, two reference strains (1330/1 Guiry 2014). A plethora of authors have suggested and 1330/2) of P. littoralis from the Culture Collection of that these various forms and varieties can possibly Algae and Protozoa (CCAP) isolated respectively in Roscoff be attributed to phenotypic plasticity (i.e., Knight in 1970 and 2002, were included in the analysis. 1923, Newton 1931, Russell 1963, 1967, Bolton DNA extraction, amplification and sequencing. DNA was € € extracted from 5 to 10 mg of dry algal tissue using the Nucle- 1979b, Muller and Stache 1989). Therefore, no â clear consensus has ever been adopted on species ospin Multi-96 plant kit (Macherey-Nagel GmbH and Co. KG, Duren,€ Germany) according to the manufacturer’s proto- delimitation within Pylaiella (Siemer and Pedersen col except for the cell lysis buffer in which samples were left 1995). Molecular tools appear as a useful alternative at room temperature for 1 h rather than heating to 65°C for to morphological observation for clarifying the sys- 30 min. Sequences of three mitochondrial genes cox1 (658 tematics of the genus Pylaiella. nt), nad1 (800 nt) and atp9 (200 nt), and of the two plastid Although various species, such as Pylaiella anglica genes rbcL (1,300 nt) and atpB (630 nt) were amplified and Ruprecht and Pylaiella aquitanica Ruprecht, have sequenced. Primers used and PCR reactions performed are described in Silberfeld et al. (2010). cox1 and rbcL genes were been reported in England and in south-western sequenced for all specimens (i.e., 140 individuals of P. littoral- France, respectively (Ruprecht 1850), Feldmann is and two CCAP strains). For the other mitochondrial and (1954), and most authors after him, recognized only chloroplast genes (nad1, atp9, and atpB), a subsample of 45 P. littoralis at the species level in Brittany, France. In individuals of P. littoralis was used to confirm results obtained this region, the species is found both on rocky with the previous genes. PCR products were purified and substrates and as an epiphyte of perennial seaweed sequenced by LGC genomics (Berlin, Germany). The species (Feldmann 1954). In the northern Baltic obtained sequences were edited and aligned using Codon- code Aligner v. 3.7 (www.codoncode.com, Centerville,
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