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Use of DNA Barcode in the Identification of Fish Species from Ribeira De Iguape Basin and Coastal Rivers from São Paulo State (Brazil)

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Use of DNA Barcode in the Identification of Fish Species from Ribeira De Iguape Basin and Coastal Rivers from São Paulo State (Brazil) DNA Barcodes 2015; Volume 3: 118–128 Research Article Open Access Jefferson Monteiro Henriques, Guilherme José da Costa Silva, Fernando Yuldi Ashikaga, Robert Hanner, Fausto Foresti and Claudio Oliveira Use of DNA barcode in the identification of fish species from Ribeira de Iguape Basin and coastal rivers from São Paulo State (Brazil) DOI 10.1515/dna-2015-0015 Received February 25, 2015; accepted July 20, 2015 1 Introduction Abstract: Species identification is a difficult task, ranging Biodiversity is threatened by problems arising from from the definition of the species concept itself to the human population growth [1,2], and species identification definition of the threshold for speciation. DNA Barcode is an important step for the preservation of biodiversity technology uses a fragment of the Cytochrome Oxidase [2,3]. However, species identification is a difficult task, I (COI) gene as a molecular tool that many studies have since many controversies have persisted to the present already validated as a tool for species identification. DNA day, ranging from the definition of the species concept barcode sequences for COI were generated and analyzed to the definition of the threshold for speciation [4]. Many from 805 specimens. The General Mixed Yule Coalescent types of methods have been proposed as the best tools for (GMYC) analysis recognized 99 independent evolution units, species identification, with genetic tools considered as an and the Barcode Index Numbers (BIN) approach pointed to alternative approach [5–12]. the existence of 104 BINs (interpreted as distinct species). By The integration of research areas for species cross-tabulating the results of all approaches, we identified identification can bring great benefits in biodiversity 109 Molecular Operational Taxonomic Units (MOTU) by characterization in regions where such information is at least one methodology. In most cases (89 MOTUs), the lacking, avoiding the loss of endemic species that may genetic approaches are in agreement with morphological perhaps occur in these regions, enabling the creation of identification, and the discrepant results of MOTUs are conservation areas. Examples of this characterization in the complex groups, which have many morphological include the checklist of ichthyofauna from the Ribeira de similarities but may represent species complexes. Iguape River Basin [13–15] and the Coastal Rivers of São Paulo [13,16]. These two water bodies are located in two Keywords: Single locus analysis, species identification of the most urbanized and densely populated regions of and delimitation, COI gene Brazil, but data on the number of freshwater fish species in both regions are incomplete. *Corresponding author: Jefferson Monteiro Henriques, Departamento Although molecular tools have provided new insights de Morfologia - Instituto de Biocięncias Universidade Estadual into several areas of study, such as evolutionary biology, Paulista - UNESP Botucatu 18618-970 Săo Paulo Brazil phylogenetic systematics, and population genetics, only Guilherme José Costa Silva, Departamento de Morfologia - Instituto in 2003 it was proposed that a short segment of 648 de Biociências Universidade Estadual Paulista - UNESP Botucatu 18618-970 São Paulo Brazil nucleotides from the mitochondrial gene cytochrome c Fernando Yuldi Ashikaga, Departamento de Morfologia - Instituto de oxidase I (COI) could standardize species identification Biociências Universidade Estadual Paulista - UNESP Botucatu 18618- [6,7]. The methodology called “DNA Barcode” 970 São Paulo Brazil subsequently has been widely used, and many studies Robert Hanner, Biodiversity Institute of Ontario and Department of Inte- have already validated it as a tool for species identification grative Biology University of Guelph Guelph N1G 2W1 Ontario Canada Fausto Foresti, Departamento de Morfologia - Instituto de Biociênci- in selected geographical areas [17–28]. as Universidade Estadual Paulista - UNESP Botucatu 18618-970 São However, many questions have been raised about the Paulo Brazil use of genetic markers for species delimitation. Population Claudio Oliveira, Departamento de Morfologia - Instituto de Biociên- genetic parameters such as gene flow and population size cias Universidade Estadual Paulista - UNESP Botucatu 18618-970 and phylogenetic parameters such as the pattern and São Paulo Brazil © 2015 Jefferson Monteiro Henriques et al. licensee De Gruyter Open. This work is licensed under the Creative Commons Attribution-NonCommercial-NoDerivs 3.0 License. Unauthenticated Download Date | 12/4/15 10:38 PM DNA barcode fish Ribeira de Iguape coastal rivers Brazil 119 timing of lineage diversification are being very actively 2 Methods studied, because the limit of genetic divergence within and between species is a factor that must be considered [25,29– 2.1 Sampling 32]. “Phylogenetic analysis is traditionally concerned with estimating relationships among species, while For this study, 805 specimens of freshwater fishes (89 population genetics is concerned with adaptation and morphospecies) were used, including 490 samples from understanding the population-level forces that change the Ribeira de Iguape Basin and 315 from Coastal Rivers allele frequencies” Carstens et al.[4]. This brings us to a (Figure 1). All specimens used were identified by specialist question: to what extent do genetic differences correspond PhD Osvaldo Takeshi Oyakawa and identification-keys. to individual variation within a population or a divergence After identification, the specimens were deposited in the between the two species? Thus, for species identification, fish collection of the Laboratory of Biology and Genetic it is necessary to use different methodologies for data of Fish (LBP), Department of Morphology, Biosciences analysis so that the conflicting data can be treated with Institute, UNESP, Botucatu, São Paulo, Brazil. (see Table S1 greater care [4]. More recent studies have used analysis of the Supplementary Material). Specimen data, including of multispecies coalescent models such as the GMYC [33] geospatial coordinates of collection sites, are recorded in for the determination of species. In addition, the Barcode the publicly accessible BOLD projects titled “Fishes from of Life Database provides a tool for analysis that allows Brazilian Coastal Rivers; Fishes from Brazilian Coastal us to identify different molecular MOTUs without prior Rivers part II” (project codes: FBCR; FBCRB), this projects morphological identification, known as the BIN [34]. were merged and analyzed as one. Based on this information, the aim of this study was to use the methodology of DNA Barcode to identify fish 2.2 DNA Extraction, PCR amplification, and species of the Ribeira de Iguape River Basin and Coastal Rivers of São Paulo, and to evaluate the efficiency of Sequencing GMYC and BIN analysis in order to determine the number DNA barcoding was performed at the Laboratório de of MOTUs in these areas. Biologia e Genética de Peixes (LBP), UNESP, Botucatu, Brazil, and at the Canadian Centre for DNA Barcoding Figure 1 Map of Ribeira de Iguape Basin (green area) and coastal rivers (red area), showing the sample sites (yellow points) where the speci- mens were obtained. Unauthenticated Download Date | 12/4/15 10:38 PM 120 J. Monteiro Henriques, et al. (CCDB), Canada. Total genomic DNA was isolated from 2.3 Sequence analysis fins or muscle tissue of each specimen using one of two different methods: with a DNeasy Tissue Kit (Qiagen), Consensus sequences from forward and reverse strands according to the manufacturer’s instructions; or with a were obtained using GENEIOUS PRO 5.4.2 [41]. Alignments vertebrate lysis buffer containing proteinase K digested were generated using MUSCLE [42] under default overnight at 56°C and subsequent extraction using a parameters. After alignment, the matrix was checked membrane-based approach on a Biomek NX (www.pall. visually for any obvious misalignments and potential com) liquid handling station using AcroPrep96 (www. cases of sequencing errors. A quality control step was beckman.com) and 1-mL filter plates with 10 mm PALL included in our workflow to detect contamination, glass fiber media [35] according to the CCDB protocol. A paralogous copies, or pseudogenes. Then, the presence of segment of approximately 648 bp from the 5ʹ end of the stop codons was checked using GENEIOUS PRO 5.4.2. mitochondrial cytochrome c oxidase subunit I (COI) gene Nucleotide variation and genetic distances were was amplified by the polymerase chain reaction (PCR) examined using MEGA 6.0 [43]. The best nucleotide using different combinations of primers: FishF1, FishR1, evolution models for the COI gene were evaluated using FishF2, FishR2 [36], the M13-tailed primer cocktails C_ MODELTEST 3.7 [44]. FishF1t1 – C_FishR1t1 and C_VF1LFt1 – C_VR1LRt1 [37], and the pair L5698-Asn [38] and H7271-COI [39]. A total volume 2.4 GMYC molecular species hypothesis of 12.5 μL of the PCR mixture included 1.25 μL of 10X buffer (10 mM Tris-HCl + 15 mM MgCl2), 0.5 μL dNTPs (200 nM The lognormal relaxed molecular clock tree was of each), 0.5 μL each 5 mM primer, 0.05 μL Platinum® estimated using BEAST v.1.6.2 [45], since GMYC requires Taq Polymerase (Invitrogen), 1 μL template DNA (12 ng), an ultrametric tree, but the tree was not time calibrated. and 8.7 μL ddH2O. The PCR reactions consisted of 30–40 The nucleotide evolutionary model used to estimate the cycles, 30 s at 95°C, 15–30 s at 48–54°C (according to ultrametric tree was the GTR+I+G model with gamma each
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