Cladocera and Anostraca from the Interior Plateau of British Columbia,Canada, as paleolimnological indicators of salinity and lake level

Cladoceran and anostracan species assemblages were identified from the surface sediments of 33 closed–basin lakes from the southern Interior Plateau of B.C. in order to explore their effectiveness as quantitative indicators of lakewater salinity and ionic composition. These lakes were chosen to maximize the range of lakewater salinity concentrations (freshwater through hypersaline) as well as brine composition (sulphate and carbonate dominated systems). The distribution of the anostracans and cladocerans were strongly correlated with lakewater salinity, ionic composition and lake depth. Based on these strong relationships significant predictive models were developed, using weighted-averaging techniques, to infer lakewater salinity based on the species composition of anostracans and cladocerans in surface sediments. Furthermore, models were developed to infer lake depth that are superior to previously used techniques based on the ratio of planktonic/littoral Cladocera. Given that the species composition of anostracans and cladocerans can be used to infer changes in salinity and lake level, and that their remains can be identified from sedimentary profiles, there is considerable potential in using their assemblages as paleolimnological indicators of past climatic conditions. This revised version was published online in July 2006 with corrections to the Cover Date.     

Late Quaternary Protoperidinium cysts as indicators of paleoproductivity in the northern Arabian Sea

The reliability of organic-walled cysts of the heterotrophic dinoflagellate Protoperidinium as paleoproductivity indicators and the influence of bottom water oxygenation on cyst preservation is assessed by using Arabian Sea records of the past 125 kyr as a natural laboratory. Multidisciplinary geochemical, micropaleontological and palynological datasets are integrated to analyze the relationship between Protoperidinium cyst concentrations and other paleoproductivity proxies. Differential preservation potential is quantified in order to establish threshold oxidative degradation values for a possible application of quantitative Protoperidinium cyst records in paleoenvironmental reconstructions. Results indicate that variations in Protoperidinium cyst concentration closely correspond to other marine productivity and/or upwelling proxies. Although oxygenation will lead to significant cyst degradation, and thus decreased concentrations, down-core patterns in Protoperidinium cyst concentration still primarily reflect changes in sea surface productivity. In view of differential preservation among dinoflagellate cysts, down-core variations in relative abundance of Protoperidinium should be treated with caution.     

Development of diatom-based salinity models for paleoclimatic research from lakes in British Columbia (Canada)

Diatoms were identified and enumerated from the surface sediments of 65 lakes located on the Cariboo and Chilcotin Plateaux (British Columbia, Canada). These lakes span a large gradient in lakewater ionic concentration (fresh through hypersaline) and composition, as well as other physical/chemical variables. Almost all of the study lakes had higher salinities in the late-summer than in the spring. The lakes with spring salinities >8 g l^–1 showed the largest seasonal increases in salinity. Ionic composition was similar in the spring and late-summer for most lakes. Both ionic concentration (i.e. salinity) and composition were important environmental variables that could account for the different diatom floras in the lakes. Diatom assemblages characteristic of carbonate-dominated and sulfate-dominated waters were identified. Other variables such as water depth and phosphorus concentration were also important.The majority (87%) of diatom taxa had estimated salinity optima < 3 g l^–1 Halophilic diatom taxa had broader tolerances to salinity when compared to the fresh water taxa, however taxa with narrow and broad tolerances could be identified across the salinity gradient. Species diversity was weakly but significantly correlated to lakewater salinity (r ² = 0.18 to 0.3, P < 0.05).Salinity inference models were developed based on the relationship between the diatom assemblages and the spring, late-summer and average salinity. The correlations between the measured and diatominferred salinity, based on the spring (r = 0.95), late-summer (r = 0.94) and average (r = 0.95) salinity data, are high because there was an extremely strong correlation (r = 0.98) between the log transformed spring and late-summer measured salinities. These salinity reconstruction models provide a tool that can be used to infer past climatic changes as part of paleolimnological studies from appropriate closed-basin lakes in British Columbia.     

Modem organic-walled dinoflagellate cysts in arctic marine environments and their (paleo-) environmental significance

The Arctic Ocean is one of the least known marine regions of the world. Because of its major influence on global climate and its hostile environmental conditions it is a fascinating area for paleoecological, paleoclimatic and paleoceanographic research. The composition of planktic microfossil assemblages, and both the trace-element and stable isotope compositions of hard parts, provide us with valuable information about the physical and biochemical parameters of surface waters in the high northern latitudes. Calcareous and biosiliceous microfossils that are traditionally used in Quaternary paleoenvironmental studies are of limited value in the Arctic Ocean because of their low abundances, low diversity and/or low preservation potential. The past several decades have seen considerable progress in our knowledge of the ecology and biogeography of dinoflagellates and their organic-walled cysts in the high northern latitudes, and these dinoflagellate cysts are now important proxies for reconstructing surface water conditions in the Quaternary. This arcticle gives an overview of the ecology of dinoflagellates and their cysts, the processes that transform the living communities into sediment communities, and the environmental gradients that may be reconstructed from fossil dinoflagellate cysts assemblages in the high northern latitudes.      

Impact of mild experimental acidification on short term invertebrate drift in a sensitive British Columbia stream

We report daytime drift behavior of lotic macroinvertebrates following short term (12 h) additions of HCl or HCl plus AlCl₃ to a circumneutral softwater (alkalinity ca. 100 µeq 1^-1) mountain stream in British Columbia, Canada. Addition of HCl (pH reduced from 7.0 to 5.9) resulted in an overall tripling of invertebrate drift density with rapid (< 1 h) increases in chironomid Diptera and Trichoptera. Small Ephemeroptera also entered the drift at high densities, but were delayed about 6 h. Addition of AlCl₃ (0.71 to 0.95 mg 1^-1 total Al³⁺) in HCl (stream pH reduced to 5.9) resulted in an overall 6-fold increase in invertebrate drift, with rapid increases by Ephemeroptera and delayed responses by chironomids and Trichoptera. These results suggest that the behavior of several macroinvertebrates from low alkalinity, unacidified streams can be altered by simulations of short-term, mild acidic deposition events. Further, the magnitude and timing of entry into the drift varies among taxonomic groups with the presence or absence of low concentrations of aluminum ions.     

Ordination of epiphytic bryophyte communities in a wet-temperate coniferous forest,South-Coastal British Columbia

Three coniferous tree species (Pseudotsuga menziesii, Tsuga heterophylla, and Thuja plicata) at Capilano Canyon, British Columbia, were studied for their epiphytic communities. Quantitative data were obtained for fourteen bryophyte species by sampling at heights of 0.5 m, 1 m and 2 m above ground level. Scapania bolanderi is an abundant and widely distributed species; Dicranum fuscescens, Bazzania denudata, Lepidozia reptans and Hypnum circinale are also common. An environmentally uniform study area was chosen to minimize the effects of factors other than bark and elevation on the distribution of epiphytes. In general, tree species are similar in terms of their epiphytic assemblages, but ordination methods using quantitative data exposed compositional variation that may be explained by differences in microclimate and bark-type. In this respect, the results point the way to further studies to examine epiphyte associations in closer detail, and to relate these associations to specific factors in the microenvironment.Nomenclature for vascular plants follows Hitchcock & Cronquist (1973); bryophyte nomenclature follows Schofield (1976) for mosses and Stotler & Crandall-Stotler (1977) for hepatics.We thank Dr. W. B. Schofield for aiding in bryophyte identification.     

Immunoglobulin (GM and KM) allotypes and relation to population history in native peoples of British Columbia: Haida and Bella Coola

Differences in the frequencies of GM haplotypes among native peoples of the Americas support the hypothesis that there were three distinct waves of migration from northeast Asia into the Americas: Paleo-Indian, Na-Dene, and Inuit (Eskimo)-Aleut (Salzano and Steinberg: Am. J. Hum. Genet. 17:273-279, 1965; Sukernik and Osipova: Hum. Genet. 61:148-153, 1982; Williams et al.:Am. J. Phys. Anthropol. 66:1-19, 1985; Szathmary: In R Kirk and E Szathmary (eds): Out of Asia: Peopling of the Americas and the Pacific. Canberra: The Journal of Pacific History, Canberra Australian National University, pp. 79-104, 1985). We studied GM allotypes in two linguistically unique populations of Canadian west coast native peoples, the Haida and the Bella Coola, and compared them to GM frequencies in populations that are supposed descendants of the three migrations, in order to investigate the possible genetic relationships of these British Columbia (BC) groups to other native populations. We also estimated the amount of European admixture from the frequency of the Caucasian haplotype, Gm3;5. Results showed that the frequencies in both BC populations of the three common native haplotypes (Gm1,17;21, Gm1,2,17;21, and Gm1,17;15,16), were intermediate between the frequencies in supposed descendants of Paleo-Indian and Na-Dene. These genetic findings are consistent with the controversial hypothesis of archeologist C. Borden (Science 203:963-971, 1979) that, following deglaciation about 13,000 years ago, British Columbia was repopulated by peoples from the north (?Na-Dene) and by culturally distinct peoples from the south (?Paleo-Indian). Caucasian admixture estimates suggested that the Haida and Bella Coola have also experienced moderate amounts (12-20%) of genetic input from European-originating peoples.(ABSTRACT TRUNCATED AT 250 WORDS)     

A life cycle evaluation of wood pellet gasification for district heating in British Columbia

The replacement of natural gas combustion for district heating by wood waste and wood pellets gasification systems with or without emission control has been investigated by a streamlined LCA. While stack emissions from controlled gasification systems are lower than the applicable regulations, compared to the current base case, 12% and 133% increases are expected in the overall human health impacts for wood pellets and wood waste, respectively. With controlled gasification, external costs and GHG emission can be reduced by 35% and 82% on average, respectively. Between wood pellets and wood waste, wood pellets appear to be the better choice as it requires less primary energy and has a much lower impact on the local air quality.     

Cribra orbitalia,a possible sign of anemia in early historic native populations of the British Columbia coast

Excavated and museum skeletons of the postcontact period revealed cribra orbitalia in four native ethnolinguistic divisions of the British Columbia coast, Haida, Kwakiutl, Nootka, and Coast Salish. Affected skulls were distributed among 25 of 35 localities, indicating widespread occurrence in a mainly heterogeneous population. Manifestations were similar to porotic hyperostosis, and additional lesions in the sample tend to support the concept that cribra orbitalia is related to anemia. Both inherited and acquired disorders may have been involved in the etiology. A uniquely high incidence of 52.9% occurred in immature of the Haida, a relatively homogeneous population. However, marked variability in expression by age group and by sex in the total sample is suggestive of iron-deficiency anemia. The data and historical information parallel modern expectations of susceptibility. Among 454 skulls, cribra orbitalia occurred in 32.7% of growing children and adolescents, 19% of infants and toddlers, 13.3% of adult females, and 4.8% of adult males. Postcontact disruptions and disease may have figured in promoting iron-deficiency anemia, but noted precontact occurrences may also have been due to the disorder.     

Climatic change and fire potential in South-Central British Columbia, Canada

The incidence and severity of forest fires are linked to the interaction between climate, fuel and topography. Increased warming and drying in the future is expected to have a significant impact on the risk of forest fire occurrence. An increase in fire risk is linked to the synchronous relationship between climate and fuel moisture conditions. A warmer, drier climate will lead to drier forest fuels that will in turn increase the chance of successful fire ignition and propagation. This interaction will increase the severity of fire weather, which, in turn, will increase the risk of extreme fire behaviour. A warmer climate will also extend fire season length, which will increase the likelihood of fires occurring over a greater proportion of the year. In this study of the North Okanagan area of British Columbia, Canada, the impacts of climate change of fire potential were evaluated using the Canadian Forest Fire Danger Rating System and multiple climate scenario analysis. Utilizing this approach, a 30% increase in fire season length was modelled to occur by 2070. In addition, statistically significant increases in fire severity and fire behaviour were also modelled. Fire weather severity was predicted to increase by 95% during the summer months by 2070 while fire behaviour was predicted to shift from surface fire‐intermittent crown fire regimes to a predominantly intermittent‐full crown fire regime by 2070 onwards. An increase in fire season length, fire weather severity and fire behaviour will increase the costs of fire suppression and the risk of property and resource loss while limiting human‐use within vulnerable forest landscapes. An increase in fire weather severity and fire behaviour over a greater proportion of the season will increase the risks faced by ecosystems and biodiversity to climatic change and increase the costs and difficulty of achieving sustainable forest management.     

Genetic diversity of Metarhizium anisopliae var. anisopliae in southwestern British Columbia

The abundance and genetic diversity of the entomopathogenic fungus, Metarhizium anisopliae var. anisopliae, in southwestern British Columbia (BC) and southern Alberta was examined. The fungus was found to be widespread in soil throughout southwestern BC, and was recovered from 56% of 85 sample sites. In contrast to southwestern BC, no M. anisopliae isolates were recovered in southern Alberta. An automated fluorescent amplified fragment length polymorphism (AFLP) method was used to examine genetic diversity. In excess of 200 isolates were characterized. The method identified 211 polymorphic amplicons, ranging in size from ≈92 to 400 base pairs, and it was found to be reproducible with a resolution limit of 86.2% similarity. The AFLP method distinguished Metarhizium from other entomopathogenic fungal genera, and demonstrated considerable genetic diversity (25 genotypes) among the reference strains of M. anisopliae isolates examined (i.e. recovered from various substrates and geographical locations). Although 13 genotypes of M. anisopliae var. anisopliae were recovered from southwestern BC soils, the vast majority of isolates (91%) belonged to one of two closely-related genotypes. Furthermore, these two genotypes predominated in urban, agricultural and forest soils. The reasons for the limited diversity of M. anisopliae var. anisopliae in southwestern BC are uncertain. However, findings of this study are consistent with island biogeography theory, and have significant implications for the development of this fungus for microbial control of pest insects.     

A molecular analysis of hybridization between native westslope cutthroat trout and introduced rainbow trout in southeastern British Columbia, Canada

Restriction site variation in the Ikaros gene intron was used to assess the incidence of westslope cutthroat trout ( Oncorhynchus clarki lewisi ), rainbow trout ( O. mykiss ) and interspecific hybrids at 11 localities among eight streams tributary to the upper Kootenay River system in south-eastern British Columbia, Canada. Out of 356 fish assayed by this technique, hybrids ( n =16) were found at seven of the 11 sites across five different streams. Rainbow trout ( n =6) were found at two of the 11 sites. Analysis of hybrids with a second genetic marker (heat shock 71 intron) indicated that most represented either backcrosses to both westslope cutthroat and rainbow trout, or post F₁ hybrids. Mitochondrial DNA analysis indicated that hybrid matings occur between male rainbow trout and female westslope cutthroat trout and vice versa. Comparison of present hybridization in five tributaries relative to an allozyme-based analysis in the mid-1980s, that documented hybrids in only a single tributary of seven that were common to the two studies, suggests that hybridization and introgression has increased in upper Kootenay River tributaries. The present analysis is a conservative estimate of genetic interaction between the species because introgression was not tested in the majority of samples. Identification of genetically pure westslope cutthroat trout populations, and why they might be resistant to introgression from rainbow trout, are crucial conservation priorities for this unique subspecies of cutthroat trout.     

Inter- and intrapopulation genetic variation in species ofPorphyra (Rhodophyta: Bangiales) from British Columbia and adjacent waters

Starch gel electrophoresis of 17 proteins has provided data on inter- and intrapopulation genetic variation in 20 species ofPorphyra occurring in British Columbia and adjacent areas.P. cuneiformis andP. nereocystis showed no within species variation, even over ranges of more than 1000 km. Populations ofP. abbottae, P. fallax, P. fucicola, P. gardneri andP. schizophylla were characterized by fixation for certain alleles. The number of polymorphic loci in a population ranged from zero to nine, depending on the species. Six species had populations that were polymorphic at just a single locus. Only two species (P. mumfordii andP. pseudolanceolata) had populations that were polymorphic at more than three loci. These levels of genetic variation are lower than those reported for populations of JapanesePorphyra species. Eleven taxa were polymorphic for 6-phosphogluconate dehydrogenase, the most variable enzyme. No within species polymorphisms were detected for bromoperoxidase, lactate dehydrogenase, superoxide dismutase or phycoerythrin. Possible evidence for the chimeric nature of the thallus was observed only inP. mumfordii.     

Calcareous dinoflagellate cysts in surface sediments from the Mediterranean Sea: distribution patterns and influence of main environmental gradients

The distribution of calcareous dinoflagellate cysts in surface sediments from the Mediterranean Sea was quantitatively analysed. The samples contain 11 cyst species and the vegetative coccoid Thoracosphaera heimii. Cyst abundance increases towards the deeper parts of the basins and is generally higher in the eastern Mediterranean Sea. Three major distribution characteristics exist: (1) different assemblages in oceanic and neritic regions, (2) little agreement with the associations of areas studied so far like the Atlantic Ocean, and (3) a unique oceanic assemblage in the eastern Mediterranean Sea. A gradual change in cyst assemblages from the western to the eastern Mediterranean Sea was observed and statistically compared with the main environmental gradients in the upper water column. Temperature, nitrate concentration and possibly salinity appear to be the most important factors controlling cyst production. Three groups containing cysts with similar environmental preferences can be distinguished: (1) an eastern Mediterranean group related to relatively high temperature and salinity but low nitrate concentration, (2) a group of more or less consistently abundant cosmopolitan species tolerating or even preferring relatively low temperature and salinity but high nitrate concentration, and (3) a group containing species that are possibly adapted to neritic environments and have probably been transported from coastal areas into the studied regions. In contrast to other calcareous plankton, calcareous dinoflagellate cysts correlate strongly with the main environmental gradients in the Mediterranean Sea, bearing a high potential for palaeoenvironmental reconstructions.     

A dendroclimatic analysis of mountain hemlock (Tsuga mertensiana) ring-width and maximum density parameters,southern British Columbia Coast Mountains,Canada

The growth of mountain hemlock trees in Pacific North America demonstrates a complex relationship to two or more seasonal environmental variables. In order to examine the radial growth response of mountain hemlock to subseasonal climate variables, ring-width and X-ray densitometric analyses were used to construct intra-annual dendroclimatic records. The intent was to highlight the difference between the dendroclimatic outcomes of standard ring-width analyses to those derived from density chronologies collected at high elevation locations in the British Columbia Coast Mountains. This study highlights the importance of using multiple tree-ring parameters to better define the complex growth behaviour in mountain hemlock trees for the construction of more robust proxy climate records. Tree-ring chronologies from three sites were used to describe the inherent climate-growth trends. Maximum tree-ring density values provided a robust data series for constructing site-specific proxy records of late-summer temperature. Annual ring-width measurements provided independent proxies of spring snowpack trends. Significant decreases in temperature and an increase in snowpack depth during the early 1700s and early 1800s coincides with documented PDO phases and Little Ice Age glacier advances. Identification of early and late growing season climate signals within mountain hemlock trees demonstrates the value of documenting the characteristics of multiple tree ring parameters in future dendroclimatic studies.     

Microsatellite DNA Population Structure and Stock Identification of Steelhead Trout (<Emphasis Type="BoldItalic">Oncorhynchus mykiss</Emphasis>) in the Nass and Skeena Rivers in Northern British Columbia

Population structure and the application to genetic stock identification for steelhead (Oncorhynchus mykiss) in the Nass and Skeena Rivers in northern British Columbia was examined using microsatellite markers. Variation at 8 microsatellite loci (Oki200, Omy77, Ots1, Ots3, Ssa85, Ots100, Ots103, and Ots108) was surveyed for approximately 930 steelhead from 7 populations in the Skeena River drainage and 850 steelhead from 10 populations in the Nass River drainage, as well as 1550 steelhead from test fisheries near the mouth of each river. Differentiation among populations within rivers accounted for about 1.9 times the variation observed among years within populations, with differences between drainages less than variation among populations within drainages. In the Nass River, winter-run populations formed a distinct group from the summer-run populations. Winter-run populations were not assessed in the Skeena River watershed. Simulated mixed-stock samples suggested that variation at the 8 microsatellite loci surveyed should provide relatively accurate and precise estimates of stock composition for fishery management applications within drainages. In the Skeena River drainage in 1998, Babine River (27%) and Bulkley drainage populations (31%) comprised the main components of the returns. For the Nass River in 1998 steelhead returning to Bell-Irving River were estimated to have comprised 39% of the fish sampled in the test fishery, with another 27% of the returns estimated to be derived from Cranberry River. The survey of microsatellite variation did not reveal enough differentiation between Nass River and Skeena River populations to be applied confidently in estimation of stock composition in marine fisheries at this time. Received January 14, 2000; accepted July 13, 2000.     

Organic-walled dinoflagellate cysts as paleoenvironmental indicators in the Paleogene; a synopsis of concepts

The organic-walled, hypnozygotic cysts of dinoflagellates (“dinocysts”) provide a rich, albeit incomplete, history of this eukaryotic plankton group in ancient sediments. Building on pioneering studies of the late 1970s and 1980s, recent ocean drilling and more detailed, integrated studies of surface sections have provided a wealth of dino-cyst data spanning the entire Paleogene. Based on multidisciplinary approaches, these studies have been instrumental in refining existing and furnishing new concepts of Paleogene paleoenvironmental and paleoclimatic reconstructions by means of dinocysts. Since dinocysts typically exhibit high abundances in neritic settings, the dinoeyst-based environmental and paleoclimatic information is complementary to that derived from typically more offshore groups such as planktonic foraminifera, coccolithophorids, diatoms, and radiolaria. While in a recent paper we gave a broad overview of case studies from around the globe (Sluijs et al. 2005), here we focus on a summary of these analyses and present a synopsis of applied paleoecological concepts in Paleogene (65–24 ma) dinocyst studies. Representing Earth’s greenhouse-icehouse transition, this episode holds the key to the understanding of extreme transient climatic change. The present paper offers guidelines for the application of dinocyst palaeoecology to the reconstruction of Paleogene sea-surface productivity, temperature, salinity, stratification, and paleo-oxygenation as well as their application in sequence stratigraphy, oceanic circulation and general watermass reconstructions.      

Size-fractionated productivity and nutrient dynamics of phytoplankton in subtropical coastal environments

It is now well established that the size distribution of phytoplankton plays an important role in primary production processes and nutrient dynamics of coastal environment. In situ observations showed that nanophytoplankton (3–20 μm) contributed 72.08% and58.18% of phytoplankton biomass and 58.32% and 41.14% of primary productivity to Xiamen Western Waters and the northern Taiwan Strait, respectively; picophytoplankton (0.2–3 μm) dominated the biomass (64.70%) and productivity (66.09%) in the southern Taiwan Strait. Furthermore, nanophytoplankton accounted for 75% of phosphate uptake with the highest rate constant (8.3×10^-5 s^-1) and uptake rate in unit water volume (5.4×10^-5 mmol dm^-3s^-1); picophytoplankton had the highest uptake rate in unit biomass (5.4×10^-5 mmol mg^-1s^-1) and photosynthetic index (3.8 mgC mgChl a^-1h^-1). All the results highlighted the remarkable characteristics of small size ranged (0.2–20 μm) phytoplankton in subtropical coastal environments: main contributor to phytoplankton biomass and production, high efficiency on organic carbon production and nutrient recycling. The far reaching environmental and ecological implications were discussed. This revised version was published online in August 2006 with corrections to the Cover Date.     

Late Holocene variability in pelagic fish scales and dinoflagellate cysts along the west coast of Vancouver Island, NE Pacific Ocean

Fish stocks and dinoflagellates are essential components of the marine food chain. Sediment cores from a predominantly anoxic basin in Effingham Inlet, Vancouver Island, British Columbia, archive a late Holocene (500–5300 years BP) record of paleoproductivity in the North American Coastal Upwelling Domain (CUD). We present evidence that late Holocene changes in the dinoflagellate cyst assemblages, sedimentary record, and fish stocks in the northeastern Pacific Ocean fluctuated, at least partially, in accordance with regional and global climate cycles.Principal components analysis (PCA), and trend, wavelet and spectral analyses were used to identify relationships, cycles and trends in sediment grey-scale values, and the abundances of fish scales and dinoflagellate cysts on centennial to millennial time scales. Most observed cycles fluctuated in intensity over time, particularly following transition of the regional climate to a higher rainfall phase that impacted coastal oceanic dynamics 3400 ± 150 years ago. Correlation of the marine paleoproductivity records observed in Effingham Inlet with solar influenced climate proxy cycles observed in the North Atlantic region indicates that solar forcing at different scales might have influenced the climate in the northeast Pacific as well. In particular an 1100- to 1400-year cycle in regional climate is well represented in the fish productivity proxy and sedimentological record. It was also observed that colder water, high-productivity, Selenopemphix nephroides and anchovy-dominated “Anchovy Regime” ecosystems alternate with warmer water, herring-dominated “Herring Regime” ecosystems at millennial time scales. The fish scale record preserved in Effingham Inlet indicates that the NE Pacific is now in transition from an ‘anchovy-’ to a ‘herring’-dominated regime.