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Lack of Genetic Differentiation of Blue Spiny Lobster Panulirus Inflatus Along the Pacific Coast of Mexico Inferred from Mtdna Sequences

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Lack of Genetic Differentiation of Blue Spiny Lobster Panulirus Inflatus Along the Pacific Coast of Mexico Inferred from Mtdna Sequences Vol. 361: 203–212, 2008 MARINE ECOLOGY PROGRESS SERIES Published June 9 doi: 10.3354/meps07381 Mar Ecol Prog Ser Lack of genetic differentiation of blue spiny lobster Panulirus inflatus along the Pacific coast of Mexico inferred from mtDNA sequences Francisco Javier García-Rodríguez1, 2, Ricardo Perez-Enriquez1,* 1Centro de Investigaciones Biológicas del Noroeste (CIBNOR), Mar Bermejo 195, Colonia Playa Palo de Santa Rita, La Paz, BCS 23090, Mexico 2Present address: Centro Interdisciplinario de Ciencias Marinas-Instituto Politécnico Nacional (CICIMAR-IPN), AP 592, La Paz, BCS 23000, Mexico ABSTRACT: Blue spiny lobster Panulirus inflatus (Bouvier, 1895) is an economically and ecologically important species along the Pacific coast of Mexico. Because most studies have been done on the local level and conducted to obtain fishery biology data, little is known about population boundaries. Previous analysis showed slight morphometric differences between lobsters along the Baja California Peninsula coast and those from the mainland coast. To determine whether differences are conse- quences of local variation or reproductive isolation and genetic variations, the genetic structure of P. inflatus was determined using sequence data of mtDNA (control region, 12S gene, and 16S gene). Genetic variability (number of haplotypes, haplotype diversity, and nucleotide diversity) for each mtDNA fragment was similar among localities. Nonsignificant differences among localities were revealed (control region: Φst = –0.0027, p = 0.555; 12S gene: Φst = –0.0011, p = 0.466; 16S gene: Φst = –0.00834, p = 0.759). Low levels of differentiation in morphology and the absence of variability of genetic structure suggest that the blue spiny lobster represents a panmictic population with pheno- typic plasticity. We suggest that the lack of variation in genetic structure is related to oceanographic processes coupled with an extended larval period. Mismatch analysis suggested that the population history is characterized by range expansion. KEY WORDS: Genetic diversity · mtDNA · Panulirus inflatus · Lobster · DNA sequencing Resale or republication not permitted without written consent of the publisher INTRODUCTION Pérez-González et al. 1992a,b, Muñoz-García et al. 2000), which has led to management measures, in- Blue spiny lobster Panulirus inflatus (Bouvier, 1895) cluding closed seasons, minimum legal size, and pro- is endemic to the tropical and subtropical Pacific coast hibition from catching egg-bearing females. More of Mexico, from southern Baja California to Oaxaca, intensive studies of population dynamics and stock including some areas in the Gulf of California (see evaluation are needed to improve the management Fig. 1), where it inhabits rocky bottoms above 30 m system (Pérez-González et al. 2002). depth (Pérez-González et al. 2002). The lobster is a Better management strategies in the exploitation of a valuable resource to the local fishery of many commu- fishery resource should be based on the recognition of nities (Vega et al. 1996). population boundaries. If the boundary of the fishery In spite of its economic importance, knowledge does not encompass a stock, adequate management about the biology of species is scarce. Information may be affected because different growth and mortal- on reproductive activity, size, and sex composition is ity characteristics of a population are not appropriately known from some studies (Briones & Lozano 1992, considered (Skillman 1989). Determination of popu- *Corresponding author. Email: [email protected] © Inter-Research 2008 · www.int-res.com 204 Mar Ecol Prog Ser 361: 203–212, 2008 lation genetic structure has not been studied in blue morphological differences in lobsters from Baja Cali- spiny lobster and only limited data are available about fornia and the mainland reflect genetic differences and disjunctive populations. García-Rodríguez et al. (2004) reproductive isolation or phenotypic plasticity associ- found significant differences in a multivariate morpho- ated with local environmental differences. metric analysis comparing the carapace of lobsters from the coast of the Baja California Peninsula and the coast of continental Mexico, suggesting that MATERIALS AND METHODS phenotypic plasticity is influenced by local differences in environmental factors rather than reproductive iso- Sampling and DNA extraction. Lobsters were lation. obtained from 5 local commercial fisheries in the Mex- Previous studies of population differences in other ican states of Baja California Sur, Sinaloa, Nayarit, lobster species were based on allozyme analyses Jalisco, and Guerrero (Fig. 1) between September (Shaklee & Samollow 1984, Hateley & Sleeter 1993, 2002 and May 2005. Lobsters from the Baja California Thompson et al. 1996, Perez-Enriquez et al. 2001). Peninsula were captured in wooden traps; at the other Recently, analysis of lineages based on mtDNA was locations, lobsters were collected with tangle nets or used to evaluate population structure in this group of diving. After collection, muscle tissue from each speci- lobsters (Diniz et al. 2005, Triantafyllidis et al. 2005, men was preserved in 70% ethanol. Total genomic García-Rodríguez & Perez-Enriquez 2006). Analysis of DNA was extracted by incubating muscle tissue mtDNA has been a useful tool to study the relationship overnight at 37°C in TNES buffer with Proteinase K between members of populations. Different mutation and using a standard phenol-chloroform procedure as rates occurring in mtDNA allow us to explore inter- described in Perez-Enriquez & Taniguchi (1999). Iso- and intraspecific genetic variations. In spiny lobster, lated DNA was suspended in 100 µl TE buffer (10 mM DNA sequencing of coding genes, such as 16S and Tris-HCl, 1 mM EDTA pH 7.6) and stored at 4°C. COI, has been used for phylogenetic reconstructions, mtDNA amplification and sequencing. PCR was supporting hypotheses based on morphological data to used to amplify 2 regions of mtDNA, a partial se- explain origins and radiation of the Panulirus genus quence of the 16S gene (543 bp) and the 12S control (George & Main 1967, Ptacek et al. 2001, George region (12S-CR) fragment, composed of a putative 2005). Sarver et al. (1998) showed that the 16S gene fragment of the 12S gene and a fragment of the control clearly helped to discriminate between 2 clades of P. region (663 bp). In both cases, each PCR reaction was argus. The control region (also known as the ‘AT-rich’ carried out in 50 µl containing PCR buffer 1X (Invitro- region) in invertebrates — which does not code for a gene), 0.2 mM dNTP mix, 0.48 µM of each primer, functional gene and therefore is under fewer func- 4.0 mM MgCl2, and 2.5 U Taq DNA polymerase (In- tional and structural restrictions — leads to high aver- vitrogene). The 12S-CR segment was amplified using age substitution rates (Saccone et al. 1987, Diniz et al. the primer pair srRNA (5’-CAGGGTATCTAATCCTG- 2005), and has also proved to be useful for population GTT; Weider et al. 1996) and PinRC1-b (5’-GATGGC- genetic studies among crustaceans (Grabowski & CCATTACCGAACTA; García-Rodriguez et al. 2008). Stuck 1999, Chu et al. 2003, McMillen- Jackson & Bert 2003, Grabowski et al. 2004, Diniz et al. 2005). Diniz et al. (2005), using the control region, showed that, in spite of the small sam- ple size, there is very high diversity and genetic structure for P. argus concor- dant with that found using more con- servative DNA sequences. Their data supports the idea that the control region has a high potential to be used in intraspecific variability studies in spiny lobsters. For the present study, we collected blue spiny lobsters at 5 locations on the Pacific coast of Mexico to study genetic structure with mtDNA sequences. We included samples from the Baja Califor- nia Peninsula to determine whether Fig. 1. Panulirus inflatus. Sampling locations along the Pacific coast of Mexico García-Rodríguez & Perez-Enriquez: Blue lobster genetics 205 PCR thermal cycling involved an initial denaturation Mismatch distribution differences were estimated to for 2 min at 94°C, followed by 35 cycles for 1 min at review historical demography. The shape of the mis- 94°C, 1 min at 63°C, and 2 min at 72°C. A final cycle, match distribution is relevant because it may be used lasting 4 min, was done at 72°C. The partial sequence to deduce whether a population is undergoing sudden of the 16S gene was amplified, using primers 16Sar-L population expansion or is in equilibrium (Rogers & and 16Sbr-H (Palumbi et al. 1991). PCR thermal condi- Harpending 1992). In general, chaotic and multimodal tions were 2 min at 94°C, 30 cycles for 1 min at 94°C, distributions suggest a population of constant size, 1 min at 60°C, and 2 min at 72°C. A final extension, whereas unimodal distributions reflect a population lasting 4 min at 72°C, was done. Amplification prod- that has experienced a sudden demographic expan- ucts underwent electrophoresis on 1% agarose gels sion. Parameters of the model of sudden expansion (τ, and stained with SybrGold, along with a molecular θ0, and θ1) and mismatch distribution differences were weight marker, to confirm amplification and expected estimated using Arlequin v. 3.0 (Excoffier et al. 2005). size. PCR products were purified and sequenced Time of expansion was estimated according to Rogers in sense and antisense (Macrogen). Individual se- & Harpending (1992): quences were deposited in GenBank (accession nos.: = τ/2µ EF645361-EF645486 for 16S gene, and EF645487- t EF645597 for the 12S-CR fragment). where t measures time in generations when a popula- Data analysis. Sequences of each gene were tion expansion begins and µ is the specific fragment arranged and overlapped using Sequencher v. 4.5 mutation rate. The parameters θ0 and θ1 measure the (GeneCode). Because of mutation rate differences expected pairwise differences before and after the between the control region and the 12S gene, the 12S- expansion. Goodness-of-fit between the observed dis- CR fragment was separated into the 12S gene and the tribution and expected distribution under the sudden control region for each sequence, based on the expansion model was computed using Arlequin sequence of Panulirus japonicus (GenBank accession (Excoffier et al.
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