Environmental factors controlling floral zonation and the distribution of burrowing and tube-dwelling organisms of Fraser delta tidal flats, British Columbia

The distribution of various burrowing and tube-dwelling organisms, their biogenic sedimentary structures, and the rates at which they turn over sediment are Investigated on three different tidal flat environments of the Fraser Delta. The organisms studied include Callianassa californiensis and Upogebia pugettensls, both thalassinldean burrowing shrimps, the burrowing polychaete Abarenicola sp., the tube-dwelling polychaetes, Praxillela sp. and Spio sp., the bivalve Mya arenaria and the gastropods Batlllaria attramentaria and Nassarius mendicus. Thalassinidean shrimps are of most interest because they are widespread over the Delta, in particular Callianassa and because their distinctive burrows are well known in the geological record. The 'marine' tidal flats of Boundary Bay on the inactive southern flank of the Fraser Delta are mantled with fine to very fine,well to very well sorted sands. The intertidal region has five floral/sedimentological zones delimited primarily by elevation and exposure and characterized by distinctive macrofaunal assemblages. These are from the shoreline seawards, the saltmarsh, algal mat, upper sand wave, eelgrass and lower sand wave zones. Topography of both small and large scale of biogenic or physical origin creates lateral heterogeneity within the biofacies of each zone. An estimated 4.25 x 10⁸ Abarenicola on Boundary Bay tidal flats annually rework about 10⁶m³ of sand. The bioturbation of this worm may be a factor limiting the extent of the algal mat zone. By irrigating its burrow, Abarenicola can separate a sand/clay mixture by floating the clay out in the head shaft irrigation current. Thalassinidean burrowing shrimps are most abundant on the 'marine' tidal flats of southeastern Roberts Bank on the active Delta-front. These tidal flats can divided into four floral/ sedimentological zones: the saltmarsh, algal mat, sandflat and eelgrass zones. Thalassinidean burrowing shrimps dominate the sandflat zone. Upogebia densities are positively correlated to mud content of the sediment. Callianassa show no clear grain size preference and are abundant in sediments ranging from 5 to 50% in mud content and from 2.6 to 4.0 Ø in median grain size. At their peak density (446 burrow openings m⁻²) Callianassa rework the substrate they live in to a depth of 50 cm in about five months. On central Roberts Bank a major transition from a 'marine' to a brackish environment occurs. A brackish marsh zone extending to much lower intertidal levels than the saltmarsh laterally replaces the algal mat zone and the upper half of the sandflat zone. A sandflat/mudflat zone cross-cut by channels displaces the eelgrass zone and lower half of the sandflat zone. The peak in Callianassa distribution moves to lower intertidal levels because of the presence of low salinity water at higher tidal levels and because of the absence of eelgrass in lower intertidal regions. Upogebia although physiologically better adapted to cope with reduced salinity demonstrates lower tolerance of brackish water in its distribution than Callianassa, probably because the function of its mud-lined burrow as a conduit for suspension feeding and respiration exposes Upogebia to low salinity surface waters, while Callianassa, in its unlined burrow used for deposit feeding, is protected from surface waters by high salinity interstitial waters. The distinction between these two types of burrow is considered to be very significant for paleoenvironmental reconstructions. A new system of subdividing the intertidal region into exposure zones (the atmozone, amphizone and aquazone), based on critical tidal levels at which the maximum duration of continuous exposure or submergence 'jumps,' is advocated. It allows cross correlation between different tidal regions experiencing different types of astronomically controlled tides and much of the intertidal zonation of Fraser Delta tidal flats may be causally related to these exposure zones.Science, Faculty ofEarth, Ocean and Atmospheric Sciences, Department ofGraduat

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Crisis and change in the system of innovation: The Japanese pharmaceutical industry during the Lost Decades, 1990–2010

This article uses the experience of the Japanese pharmaceutical industry to show how Japan's national system of innovation evolved from a closed, firm-based domestic system toward a more open, networked, global system. This occurred in the face of a crisis of economic and technological dimensions. During the Lost Decades, the nature of innovation in this industry shifted from incremental toward more radical innovation, as the system internationalised and as firms leveraged different environments around the world. This article highlights the varying roles that the components of the system of innovation play in shaping innovative industries. It also shows how institutions can be remarkably malleable in times of crisis