965 resultados para Thermal structure in the sea
Resumo:
Data on zooplankton abundance and biovolume were collected in concert with data on the biophysical environment at 9 stations in the North Atlantic, from the Iceland Basin in the East to the Labrador Sea in the West. The data were sampled along vertical profiles by a Laser Optical Plankton Counter (LOPC, Rolls Royce Canada Ltd.) that was mounted on a carousel water sampler together with a Conductivity-Temperature-Depth sensor (CTD, SBE19plusV2, Seabird Electronics, Inc., USA) and a fluorescence sensor (F, ECO Puck chlorophyll a fluorometer, WET Labs Inc., USA). Based on the LOPC data, abundance (individuals/m**3) and biovolume (mm3/m**3) were calculated as described in the LOPC Software Operation Manual [(Anonymous, 2006), http://www.brooke-ocean.com/index.html]. LOPC data were regrouped into 49 size groups of equal log10(body volume) increments, see Edvardsen et al. (2002, doi:10.3354/meps227205). LOPC data quality was checked as described in Basedow et al. (2013, doi:10.1016/j.pocean.2012.10.005). Fluorescence was roughly converted into chlorophyll based on filtered chlorophyll values obtained from station 10 in the Labrador Sea. Due to the low number of filtered samples that was used for the conversion the resulting chlorophyll values should be considered with care. CTD data were screened for erroneous (out of range) values and then averaged to the same frequency as the LOPC data (2 Hz). All data were processed using especially developed scripts in the python programming language. The LOPC is an optical instrument designed to count and measure particles (0.1 to 30 mm equivalent spherical diameter) in the water column, see Herman et al., (2004, doi:10.1093/plankt/fbh095). The size of particles as equivalent spherical diameter (ESD) was computed as described in the manual (Anonymous, 2006), and in more detail in Checkley et al. (2008, doi:10.4319/lo.2008.53.5_part_2.2123) and Gaardsted et al. (2010, doi:10.1111/j.1365-2419.2010.00558.x).
Resumo:
Data on zooplankton abundance and biovolume were collected in concert with data on the biophysical environment at 9 stations in the North Atlantic, from the Iceland Basin in the East to the Labrador Sea in the West. The data were sampled along vertical profiles by a Laser Optical Plankton Counter (LOPC, Rolls Royce Canada Ltd.) that was mounted on a carousel water sampler together with a Conductivity-Temperature-Depth sensor (CTD, SBE19plusV2, Seabird Electronics, Inc., USA) and a fluorescence sensor (F, ECO Puck chlorophyll a fluorometer, WET Labs Inc., USA). Based on the LOPC data, abundance (individuals/m**3) and biovolume (mm3/m**3) were calculated as described in the LOPC Software Operation Manual [(Anonymous, 2006), http://www.brooke-ocean.com/index.html]. LOPC data were regrouped into 49 size groups of equal log10(body volume) increments, see Edvardsen et al. (2002, doi:10.3354/meps227205). LOPC data quality was checked as described in Basedow et al. (2013, doi:10.1016/j.pocean.2012.10.005). Fluorescence was roughly converted into chlorophyll based on filtered chlorophyll values obtained from station 10 in the Labrador Sea. Due to the low number of filtered samples that was used for the conversion the resulting chlorophyll values should be considered with care. CTD data were screened for erroneous (out of range) values and then averaged to the same frequency as the LOPC data (2 Hz). All data were processed using especially developed scripts in the python programming language. The LOPC is an optical instrument designed to count and measure particles (0.1 to 30 mm equivalent spherical diameter) in the water column, see Herman et al., (2004, doi:10.1093/plankt/fbh095). The size of particles as equivalent spherical diameter (ESD) was computed as described in the manual (Anonymous, 2006), and in more detail in Checkley et al. (2008, doi:10.4319/lo.2008.53.5_part_2.2123) and Gaardsted et al. (2010, doi:10.1111/j.1365-2419.2010.00558.x).
Resumo:
Data on zooplankton abundance and biovolume were collected in concert with data on the biophysical environment at 9 stations in the North Atlantic, from the Iceland Basin in the East to the Labrador Sea in the West. The data were sampled along vertical profiles by a Laser Optical Plankton Counter (LOPC, Rolls Royce Canada Ltd.) that was mounted on a carousel water sampler together with a Conductivity-Temperature-Depth sensor (CTD, SBE19plusV2, Seabird Electronics, Inc., USA) and a fluorescence sensor (F, ECO Puck chlorophyll a fluorometer, WET Labs Inc., USA). Based on the LOPC data, abundance (individuals/m**3) and biovolume (mm3/m**3) were calculated as described in the LOPC Software Operation Manual [(Anonymous, 2006), http://www.brooke-ocean.com/index.html]. LOPC data were regrouped into 49 size groups of equal log10(body volume) increments, see Edvardsen et al. (2002, doi:10.3354/meps227205). LOPC data quality was checked as described in Basedow et al. (2013, doi:10.1016/j.pocean.2012.10.005). Fluorescence was roughly converted into chlorophyll based on filtered chlorophyll values obtained from station 10 in the Labrador Sea. Due to the low number of filtered samples that was used for the conversion the resulting chlorophyll values should be considered with care. CTD data were screened for erroneous (out of range) values and then averaged to the same frequency as the LOPC data (2 Hz). All data were processed using especially developed scripts in the python programming language. The LOPC is an optical instrument designed to count and measure particles (0.1 to 30 mm equivalent spherical diameter) in the water column, see Herman et al., (2004, doi:10.1093/plankt/fbh095). The size of particles as equivalent spherical diameter (ESD) was computed as described in the manual (Anonymous, 2006), and in more detail in Checkley et al. (2008, doi:10.4319/lo.2008.53.5_part_2.2123) and Gaardsted et al. (2010, doi:10.1111/j.1365-2419.2010.00558.x).
Resumo:
Data on zooplankton abundance and biovolume were collected in concert with data on the biophysical environment at 9 stations in the North Atlantic, from the Iceland Basin in the East to the Labrador Sea in the West. The data were sampled along vertical profiles by a Laser Optical Plankton Counter (LOPC, Rolls Royce Canada Ltd.) that was mounted on a carousel water sampler together with a Conductivity-Temperature-Depth sensor (CTD, SBE19plusV2, Seabird Electronics, Inc., USA) and a fluorescence sensor (F, ECO Puck chlorophyll a fluorometer, WET Labs Inc., USA). Based on the LOPC data, abundance (individuals/m**3) and biovolume (mm3/m**3) were calculated as described in the LOPC Software Operation Manual [(Anonymous, 2006), http://www.brooke-ocean.com/index.html]. LOPC data were regrouped into 49 size groups of equal log10(body volume) increments, see Edvardsen et al. (2002, doi:10.3354/meps227205). LOPC data quality was checked as described in Basedow et al. (2013, doi:10.1016/j.pocean.2012.10.005). Fluorescence was roughly converted into chlorophyll based on filtered chlorophyll values obtained from station 10 in the Labrador Sea. Due to the low number of filtered samples that was used for the conversion the resulting chlorophyll values should be considered with care. CTD data were screened for erroneous (out of range) values and then averaged to the same frequency as the LOPC data (2 Hz). All data were processed using especially developed scripts in the python programming language. The LOPC is an optical instrument designed to count and measure particles (0.1 to 30 mm equivalent spherical diameter) in the water column, see Herman et al., (2004, doi:10.1093/plankt/fbh095). The size of particles as equivalent spherical diameter (ESD) was computed as described in the manual (Anonymous, 2006), and in more detail in Checkley et al. (2008, doi:10.4319/lo.2008.53.5_part_2.2123) and Gaardsted et al. (2010, doi:10.1111/j.1365-2419.2010.00558.x).
Resumo:
Data on zooplankton abundance and biovolume were collected in concert with data on the biophysical environment at 9 stations in the North Atlantic, from the Iceland Basin in the East to the Labrador Sea in the West. The data were sampled along vertical profiles by a Laser Optical Plankton Counter (LOPC, Rolls Royce Canada Ltd.) that was mounted on a carousel water sampler together with a Conductivity-Temperature-Depth sensor (CTD, SBE19plusV2, Seabird Electronics, Inc., USA) and a fluorescence sensor (F, ECO Puck chlorophyll a fluorometer, WET Labs Inc., USA). Based on the LOPC data, abundance (individuals/m**3) and biovolume (mm3/m**3) were calculated as described in the LOPC Software Operation Manual [(Anonymous, 2006), http://www.brooke-ocean.com/index.html]. LOPC data were regrouped into 49 size groups of equal log10(body volume) increments, see Edvardsen et al. (2002, doi:10.3354/meps227205). LOPC data quality was checked as described in Basedow et al. (2013, doi:10.1016/j.pocean.2012.10.005). Fluorescence was roughly converted into chlorophyll based on filtered chlorophyll values obtained from station 10 in the Labrador Sea. Due to the low number of filtered samples that was used for the conversion the resulting chlorophyll values should be considered with care. CTD data were screened for erroneous (out of range) values and then averaged to the same frequency as the LOPC data (2 Hz). All data were processed using especially developed scripts in the python programming language. The LOPC is an optical instrument designed to count and measure particles (0.1 to 30 mm equivalent spherical diameter) in the water column, see Herman et al., (2004, doi:10.1093/plankt/fbh095). The size of particles as equivalent spherical diameter (ESD) was computed as described in the manual (Anonymous, 2006), and in more detail in Checkley et al. (2008, doi:10.4319/lo.2008.53.5_part_2.2123) and Gaardsted et al. (2010, doi:10.1111/j.1365-2419.2010.00558.x).
Resumo:
Data on zooplankton abundance and biovolume were collected in concert with data on the biophysical environment at 9 stations in the North Atlantic, from the Iceland Basin in the East to the Labrador Sea in the West. The data were sampled along vertical profiles by a Laser Optical Plankton Counter (LOPC, Rolls Royce Canada Ltd.) that was mounted on a carousel water sampler together with a Conductivity-Temperature-Depth sensor (CTD, SBE19plusV2, Seabird Electronics, Inc., USA) and a fluorescence sensor (F, ECO Puck chlorophyll a fluorometer, WET Labs Inc., USA). Based on the LOPC data, abundance (individuals/m**3) and biovolume (mm3/m**3) were calculated as described in the LOPC Software Operation Manual [(Anonymous, 2006), http://www.brooke-ocean.com/index.html]. LOPC data were regrouped into 49 size groups of equal log10(body volume) increments, see Edvardsen et al. (2002, doi:10.3354/meps227205). LOPC data quality was checked as described in Basedow et al. (2013, doi:10.1016/j.pocean.2012.10.005). Fluorescence was roughly converted into chlorophyll based on filtered chlorophyll values obtained from station 10 in the Labrador Sea. Due to the low number of filtered samples that was used for the conversion the resulting chlorophyll values should be considered with care. CTD data were screened for erroneous (out of range) values and then averaged to the same frequency as the LOPC data (2 Hz). All data were processed using especially developed scripts in the python programming language. The LOPC is an optical instrument designed to count and measure particles (0.1 to 30 mm equivalent spherical diameter) in the water column, see Herman et al., (2004, doi:10.1093/plankt/fbh095). The size of particles as equivalent spherical diameter (ESD) was computed as described in the manual (Anonymous, 2006), and in more detail in Checkley et al. (2008, doi:10.4319/lo.2008.53.5_part_2.2123) and Gaardsted et al. (2010, doi:10.1111/j.1365-2419.2010.00558.x).
Resumo:
Data on zooplankton abundance and biovolume were collected in concert with data on the biophysical environment at 9 stations in the North Atlantic, from the Iceland Basin in the East to the Labrador Sea in the West. The data were sampled along vertical profiles by a Laser Optical Plankton Counter (LOPC, Rolls Royce Canada Ltd.) that was mounted on a carousel water sampler together with a Conductivity-Temperature-Depth sensor (CTD, SBE19plusV2, Seabird Electronics, Inc., USA) and a fluorescence sensor (F, ECO Puck chlorophyll a fluorometer, WET Labs Inc., USA). Based on the LOPC data, abundance (individuals/m**3) and biovolume (mm3/m**3) were calculated as described in the LOPC Software Operation Manual [(Anonymous, 2006), http://www.brooke-ocean.com/index.html]. LOPC data were regrouped into 49 size groups of equal log10(body volume) increments, see Edvardsen et al. (2002, doi:10.3354/meps227205). LOPC data quality was checked as described in Basedow et al. (2013, doi:10.1016/j.pocean.2012.10.005). Fluorescence was roughly converted into chlorophyll based on filtered chlorophyll values obtained from station 10 in the Labrador Sea. Due to the low number of filtered samples that was used for the conversion the resulting chlorophyll values should be considered with care. CTD data were screened for erroneous (out of range) values and then averaged to the same frequency as the LOPC data (2 Hz). All data were processed using especially developed scripts in the python programming language. The LOPC is an optical instrument designed to count and measure particles (0.1 to 30 mm equivalent spherical diameter) in the water column, see Herman et al., (2004, doi:10.1093/plankt/fbh095). The size of particles as equivalent spherical diameter (ESD) was computed as described in the manual (Anonymous, 2006), and in more detail in Checkley et al. (2008, doi:10.4319/lo.2008.53.5_part_2.2123) and Gaardsted et al. (2010, doi:10.1111/j.1365-2419.2010.00558.x).
Resumo:
Data on zooplankton abundance and biovolume were collected in concert with data on the biophysical environment at 9 stations in the North Atlantic, from the Iceland Basin in the East to the Labrador Sea in the West. The data were sampled along vertical profiles by a Laser Optical Plankton Counter (LOPC, Rolls Royce Canada Ltd.) that was mounted on a carousel water sampler together with a Conductivity-Temperature-Depth sensor (CTD, SBE19plusV2, Seabird Electronics, Inc., USA) and a fluorescence sensor (F, ECO Puck chlorophyll a fluorometer, WET Labs Inc., USA). Based on the LOPC data, abundance (individuals/m**3) and biovolume (mm3/m**3) were calculated as described in the LOPC Software Operation Manual [(Anonymous, 2006), http://www.brooke-ocean.com/index.html]. LOPC data were regrouped into 49 size groups of equal log10(body volume) increments, see Edvardsen et al. (2002, doi:10.3354/meps227205). LOPC data quality was checked as described in Basedow et al. (2013, doi:10.1016/j.pocean.2012.10.005). Fluorescence was roughly converted into chlorophyll based on filtered chlorophyll values obtained from station 10 in the Labrador Sea. Due to the low number of filtered samples that was used for the conversion the resulting chlorophyll values should be considered with care. CTD data were screened for erroneous (out of range) values and then averaged to the same frequency as the LOPC data (2 Hz). All data were processed using especially developed scripts in the python programming language. The LOPC is an optical instrument designed to count and measure particles (0.1 to 30 mm equivalent spherical diameter) in the water column, see Herman et al., (2004, doi:10.1093/plankt/fbh095). The size of particles as equivalent spherical diameter (ESD) was computed as described in the manual (Anonymous, 2006), and in more detail in Checkley et al. (2008, doi:10.4319/lo.2008.53.5_part_2.2123) and Gaardsted et al. (2010, doi:10.1111/j.1365-2419.2010.00558.x).
Resumo:
Data on zooplankton abundance and biovolume were collected in concert with data on the biophysical environment at 9 stations in the North Atlantic, from the Iceland Basin in the East to the Labrador Sea in the West. The data were sampled along vertical profiles by a Laser Optical Plankton Counter (LOPC, Rolls Royce Canada Ltd.) that was mounted on a carousel water sampler together with a Conductivity-Temperature-Depth sensor (CTD, SBE19plusV2, Seabird Electronics, Inc., USA) and a fluorescence sensor (F, ECO Puck chlorophyll a fluorometer, WET Labs Inc., USA). Based on the LOPC data, abundance (individuals/m**3) and biovolume (mm3/m**3) were calculated as described in the LOPC Software Operation Manual [(Anonymous, 2006), http://www.brooke-ocean.com/index.html]. LOPC data were regrouped into 49 size groups of equal log10(body volume) increments, see Edvardsen et al. (2002, doi:10.3354/meps227205). LOPC data quality was checked as described in Basedow et al. (2013, doi:10.1016/j.pocean.2012.10.005). Fluorescence was roughly converted into chlorophyll based on filtered chlorophyll values obtained from station 10 in the Labrador Sea. Due to the low number of filtered samples that was used for the conversion the resulting chlorophyll values should be considered with care. CTD data were screened for erroneous (out of range) values and then averaged to the same frequency as the LOPC data (2 Hz). All data were processed using especially developed scripts in the python programming language. The LOPC is an optical instrument designed to count and measure particles (0.1 to 30 mm equivalent spherical diameter) in the water column, see Herman et al., (2004, doi:10.1093/plankt/fbh095). The size of particles as equivalent spherical diameter (ESD) was computed as described in the manual (Anonymous, 2006), and in more detail in Checkley et al. (2008, doi:10.4319/lo.2008.53.5_part_2.2123) and Gaardsted et al. (2010, doi:10.1111/j.1365-2419.2010.00558.x).
Resumo:
The objective of this study was to test whether elevated pCO2 predicted for the year 2100 (85.1 Pa) affects bleaching in the coral Seriatopora caliendrum (Ehrenberg 1834) either independently or interactively with high temperature (30.5 °C). Response variables detected the sequence of events associated with the onset of bleaching: reduction in the photosynthetic performance of symbionts as measured by maximum photochemical efficiency (F v/F m) and effective photochemical efficiency (delta F/F m') of PSII, declines in net photosynthesis (P net) and photosynthetic efficiency (alpha), and finally, reduced chlorophyll a and symbiont concentrations. S. caliendrum was collected from Nanwan Bay, Taiwan, and subjected to combinations of temperature (27.7 vs. 30.5 °C) and pCO2 (45.1 vs. 85.1 Pa) for 14 days. High temperature reduced values of all dependent variables (i.e., bleaching occurred), but high pCO2 did not affect Symbiodinium photophysiology or productivity, and did not cause bleaching. These results suggest that short-term exposure to 81.5 Pa pCO2, alone and in combination with elevated temperature, does not cause or affect coral bleaching.
Resumo:
Tese de doutoramento, Biologia (Biologia Marinha e Aquacultura), Universidade de Lisboa, Faculdade de Ciências, 2016
Resumo:
Lake Annie is a small (37 ha), relatively deep (21 m) sinkhole lake on the Lake Wales Ridge (LWR) of central Florida with a long history of study, including monthly limnological monitoring since June, 1983. The record shows high variability in Secchi disc transparency, which ranged from < 1 to 15 m with a trend toward decreasing values over the latter decade of record. We examined available regional meteorological, groundwater and limnological data to determine the drivers and thermal consequences of variability in water transparency. While total nutrient concentrations and chlorophyll-a were highest during years of low transparency, stepwise regression showed that none of these had a signifi cant effect on transparency after water color was taken into account. Repeated years of high precipitation between 1993–2005 caused an increase in water table height, increasing the transport of dissolved substances from the vegetated watershed into the lake. Groundwater stage explained 73 % of the interannual variability in water transparency. Transparency, in turn, explained 85 % of the interannual variability in the heat budget for the lake, which ranged from 1.8 × 108 to 4.1 × 108 Joules m–2 yr–1, encompassing the range reported across Florida lakes. While surface water temperature was not affected by transparency, depths below 5 m warmed faster during the stratifi ed period during years having a lower rate of light extinction. We show that an increase in precipitation of 20 cm per year reduces the depth of the summer euphotic zone and thermocline by 1.9 and 1.6 m, respectively, and causes a 1-month reduction in the duration of winter mixing in this monomictic lake. Because biota have been shown to respond to shifts in light and heat distribution of much smaller magnitude than exhibited here, our work suggests that subtle changes in precipitation linked to climate fl uctuations may have signifi cant physical as well as biotic consequences.
Resumo:
Lampreys are a group of ancient vertebrates with 360 million years of existence. Throughout their evolution, they have acquired local adaptations to the colonized habitats, showing high plasticity and adaptive capacities. The sea lamprey (Petromyzon marinus L.) is a parasitic and anadromous species that occurs in both sides of the North Atlantic. The aims of this study were to analyse, using microsatellite markers, the genetic diversity and population structure of sea lamprey throughout its distributional range. Analyses demonstrated consistent signs of high population differentiation between European and North American samples (two-groups structure), most probably due to isolation by distance, but low differentiation among populations from the same coast. The apparent lack of homing in this species is in line with its high evolutive success, as homing may bring adults back to natal habitats that have changed, or that are intermittently unfavourable. Analyses also demonstrated higher levels of genetic diversity in North American samples; DIVERSIDADE GENÉTICA E ESTRUTURA POPULACIONAL DA LAMPREIA-MARINHA (PETROMYZON MARINUS L.) AO LONGO DA SUA ÁREA DE DISTRIBUIÇÃO Resumo: As lampreias são organismos ancestrais com cerca de 360 milhões de anos de existência. No decorrer da longa escala evolutiva têm vindo a adquirir adaptações aos locais que colonizaram, tendo uma forte capacidade evolutiva e adaptativa. A lampreia-marinha (Petromyzon marinus L.) é uma espécie parasita e anádroma que ocorre em ambas as costas do Atlântico Norte. Este estudo teve como principal objetivo estudar a diversidade genética e a estrutura populacional desta espécie ao longo da sua área de distribuição, através do uso de microssatélites. Os resultados demonstraram forte divergência entre populações das costas Este e Oeste do Atlântico Norte, provavelmente devido à elevada distância entre populações, mas pouca diferenciação entre populações da mesma costa. A ausência de homing nesta espécie terá contribuído para o seu sucesso evolutivo, uma vez que o homing pode levar indivíduos a reproduzirem-se em habitats que se tornaram desfavoráveis ou intermitentemente inapropriados. Os resultados demonstraram também uma maior variabilidade genética nas populações americanas.
Resumo:
Structure of intertidal and subtidal benthic macrofauna in the northeastern region of Todos os Santos Bay (TSB), northeast Brazil, was investigated during a period of two years. Relationships with environmental parameters were studied through uni-and multivariate statistical analyses, and the main distributional patterns shown to be especially related to sediment type and content of organic fractions (Carbon, Nitrogen, Phosphorus), on both temporal and spatial scales. Polychaete annelids accounted for more than 70% of the total fauna and showed low densities, species richness and diversity, except for the area situated on the reef banks. These banks constitute a peculiar environment in relation to the rest of the region by having coarse sediments poor in organic matter and rich in biodetritic carbonates besides an abundant and diverse fauna. The intertidal region and the shallower area nearer to the oil refinery RLAM, with sediments composed mainly of fine sand, seem to constitute an unstable system with few highly dominant species, such as Armandia polyophthalma and Laeonereis acuta. In the other regions of TSB, where muddy bottoms predominated, densities and diversity were low, especially in the stations near the refinery. Here the lowest values of the biological indicators occurred together with the highest organic compound content. In addition, the nearest sites (stations 4 and 7) were sometimes azoic. The adjacent Caboto, considered as a control area at first, presented low density but intermediate values of species diversity, which indicates a less disturbed environment in relation to the pelitic infralittoral in front of the refinery. The results of the ordination analyses evidenced five homogeneous groups of stations (intertidal; reef banks; pelitic infralittoral; mixed sediments; Caboto) with different specific patterns, a fact which seems to be mainly related to granulometry and chemical sediment characteristics.
Resumo:
Background: The ideal malaria parasite populations for initial mapping of genomic regions contributing to phenotypes such as drug resistance and virulence, through genome-wide association studies, are those with high genetic diversity, allowing for numerous informative markers, and rare meiotic recombination, allowing for strong linkage disequilibrium (LD) between markers and phenotype-determining loci. However, levels of genetic diversity and LD in field populations of the major human malaria parasite P. vivax remain little characterized. Results: We examined single-nucleotide polymorphisms (SNPs) and LD patterns across a 100-kb chromosome segment of P. vivax in 238 field isolates from areas of low to moderate malaria endemicity in South America and Asia, where LD tends to be more extensive than in holoendemic populations, and in two monkey-adapted strains (Salvador-I, from El Salvador, and Belem, from Brazil). We found varying levels of SNP diversity and LD across populations, with the highest diversity and strongest LD in the area of lowest malaria transmission. We found several clusters of contiguous markers with rare meiotic recombination and characterized a relatively conserved haplotype structure among populations, suggesting the existence of recombination hotspots in the genome region analyzed. Both silent and nonsynonymous SNPs revealed substantial between-population differentiation, which accounted for similar to 40% of the overall genetic diversity observed. Although parasites clustered according to their continental origin, we found evidence for substructure within the Brazilian population of P. vivax. We also explored between-population differentiation patterns revealed by loci putatively affected by natural selection and found marked geographic variation in frequencies of nucleotide substitutions at the pvmdr-1 locus, putatively associated with drug resistance. Conclusion: These findings support the feasibility of genome-wide association studies in carefully selected populations of P. vivax, using relatively low densities of markers, but underscore the risk of false positives caused by population structure at both local and regional levels.