Use of substratum ripples for flow refuging by Atlantic cod, Gadus morhua

The ability to maintain position in a current without actively swimming (station-holding) was measured on substratum ripples for Atlantic cod, Gadus morhua, a bentho-pelagic fusiform species. The current velocities tested ranged from 0–111 cm sec^-1. Ripples were sinusoidal, with twelve combinations of ripple wavelength (10, 25, 50, 125 cm) and ripple amplitude (1.0, 2.5, 5.0 cm). Ripple wavelengths were chosen to approximate 0.5, 1.0, 2.0 and 5.0 times fish total length. The potential of ripples to locally retard current and thereby provide a refuge from the flow was measured as a velocity ratio, u_trough/u_free-stream, where u_trough is the flow velocity measured at a height of 0.5 cm from the bottom of a trough and u_free-stream the flow velocity measured at a height of 10 cm above ripple crests. Cod usually swam steadily above substratum ripple crests in the free-stream flow. They used substratum ripples to hold station on only 3 of the 12 ripples tested by refuging from the flow in the ripple troughs (flow refuging). These ripples had wavelengths approaching twice the body length, with ripple amplitudes that produced velocity ratios of 0.44–0.65, thus providing at least a 35% flow reduction in the troughs. In addition, these ripples were only used at intermediate velocities starting at 49–78 cm sec^-1 and ending at 81–109 cm sec^-1 depending on the ripple morphology, suggesting there may be costs involved in flow refuging, probably in stability control. Flow refuging on substratum ripples could dramatically impact the physiology and ecology of cod in high current velocities by providing areas of retreat for energetic savings, but also offering opportunities for enhanced feeding and migration.     

Microhabitat preferences of benthic fauna in a woodland stream

Estimates of numbers, biomass, and diversity of benthic macroinvertebrates were made quarterly over a two-year period to investigate microhabitat preferences. Although biomass of most taxa was significantly different among sampling times, physical factors also appeared to be important in determining abundance of many taxa. Optimum depth, velocity, substrate type, and turbulence were determined for major taxa. Optimum conditions for diversity appeared to be 34 cm depth, 60 cm s^?1 velocity, and rubble and boulder substrate type. Habitat preference functions were derived for several taxa based on significant polynomial regressions of biomass on depth, velocity, substrate, and Froude number (turbulence). The relationship between abundance and physical habitat conditions was tested by using the product of the preference factors (range: 0–1) for depth, velocity and substrate type as a measure of habitat suitability (joint preference factor). There were significant correlations between biomass [transformed by log_e (x + 1)] of 10 benthic species and the joint preference factor. The joint preference factors accounted for from 11 to 61% of the variation of biomass of the 10 benthic species. The intercepts of the relationships between biomass of individual species and the joint preference factor were not significantly different from zero for any species. Therefore, the joint preference factors appear to be valid indicators of biomass. The preference functions have utility in habitat assessment studies, specifically with regard to minimum instream flow determinations.     

Ion and sugar permeabilities of lecithin bilayers: Comparison of curved and planar bilayers

Summary Na⁺ and sugar permeabilities of egg lecithin bilayers were measured using curved bilayers and planar bilayers as represented by single-bilayer vesicles and black lipid films, respectively. The Na⁺ permeability coefficient measured with single-bilayer vesicles at 25°C is (2.1±0.6)×10^–13 cm sec^–1. Because of technical difficulties it has been impossible to measure ionic permeabilities of values lower than about 10^–10 cm sec^–1 in planar (black) lipid bilayers using tracer methods. Thed-glucose andd-fructose permeabilities were measured with both curved and planar bilayers. The permeability coefficients measured with vesicles at 25°C are (0.3±0.2)×10^–10 cm sec^–1 for glucose and (4±1)×10^–10 cm sec^–1 ford-fructose; these are in reasonable agreement with the corresponding values obtained for planar (black) lipid bilayers which are (1.1±0.3)×10^–10 cm sec^–1 ford-glucose and (9.3±0.3)×10^–10 cm sec^–1 ford-fructose, respectively.This paper is dedicated to the memory of Walther Wilbrandt,cuius nomini nullum par elogium.     

Effects of copper on algal communities at different current velocities

This study examines the influence of current velocity in the toxiceffect of copper in diatom-dominated biofilms grown in artificial channels.Effects on community structure, algal biomass and photosynthesis (carbonincorporation) caused by 15 g L^–1 of copperwere tested at contrasting (1 and 15 cm s^–1)velocities. Moreover, a possible threshold on the effect of copper on algalbiomass and photosynthesis related to current velocity was examined by usingprogressively increasing current velocity (1 to 50 cms^–1) at 15 g L^–1 Cu.Chlorophyll-a decreased ca. 50% as a result of addition of15 g L^–1 Cu. Chlorophyll decrease occurredearlier at 15 cm s^–1 than at 1 cms^–1 when adding 15 g L^–1Cu. Copper also caused a remarkable decrease in carbon incorporation(from 30 to ca. 50%), which was produced earlier at 15 cms^–1 (three days) than at 1 cms^–1 (seven days). Some taxa were affected by thecombination of copper and current velocity. Both Achnanthesminutissima and Stigeoclonium tenue becomedominant at 15 cm s^–1 in the presence of copper.Significant inhibition of algal growth in 15 g L^–1Cu occurred at low (1 cm s^–1) and highvelocities (50 cm s^–1), but not at intermediatevelocity (20 cm s^–1). The experiments indicatethat current velocity triggers the effect that copper has on diatom-dominatedbiofilms, and that the effect is more remarkable at low and high than atintermediate current velocities.     

Dynamics of intertidal marshes near shallow estuaries in Louisiana

A three-year (1991–1993) field investigation was conducted to quantify the hydrodynamics of intertidal marshes adjacent to tidal channels and shallow bays within two Louisiana coastal regions: (1) the sediment-rich Atchafalaya Basin, and, (2) the sediment-poor Terrebonne Basin with relatively minor riverine inflow. The Terrebonne Basin marsh is regularly inundated and flooding is characterized by sporadic draining interspersed by prolonged flooding events. The maximum water depth on the marsh surface exceeds 50 cm, the flow velocity across marsh surface reaches 10 cm sec^–1, and the sediment deposition rate varies from 10 to 90 g m^–2 per tidal cycle. This rather high sediment deposition rate occurs during winter storms with strong southerly winds. In contrast, the marsh site within the sediment-rich Atchafalaya Basin is irregularly inundated and characterized by sporadic flooding interspersed by prolonged draining. There the marsh flooding depth rarely exceeds 25 cm, the over-marsh flow velocity barely reaches 2.5 cm sec^–1, and the sediment deposition rate ranges from 5 to 50 g m^–2 per tidal cycle. The surprisingly low rate of sediment deposition in a marsh within a sediment-rich region is largely due to the man-made canals that alter the hydrologic regime in the upper reaches of the tidal channel.     

Depth distribution of microbial production and oxidation of methane in northern boreal peatlands

The depth distributions of anaerobic microbial methane production and potential aerobic microbial methane oxidation were assessed at several sites in both Sphagnum- and sedge-dominated boreal peatlands in Sweden, and compared with net methane emissions from the same sites. Production and oxidation of methane were measured in peat slurries, and emissions were measured with the closed-chamber technique. Over all eleven sites sampled, production was, on average, highest 12 cm below the depth of the average water table. On the other hand, highest potential oxidation of methane coincided with the depth of the average water table. The integrated production rate in the 0–60 cm interval ranged between 0.05 and 1.7 g CH4 m ^–2 day^– and was negatively correlated with the depth of the average water table (linear regression: r ² = 0.50, P = 0.015). The depth-integrated potential CH₄-oxidation rate ranged between 3.0 and 22.1 g CH₄ m^–2 day^–1 and was unrelated to the depth of the average water table. A larger fraction of the methane was oxidized at sites with low average water tables; hence, our results show that low net emission rates in these environments are caused not only by lower methane production rates, but also by conditions more favorable for the development of CH₄-oxidizing bacteria in these environments. Correspondence to: I. Sundh.     

Photosynthesis and successional status of Costa Rican rain forest trees

Seven tree species from three different light environments in the wet lowland forests of Costa Rica were grown under controlled environment conditions to assess light related photosynthetic potentials. Light saturated photosynthesis rates were clearly related to light levels of the field environments. Mean saturated, net photosynthetic rates ranged from 6.8 to 11.3 to 27.7 mol m^–2 sec^–1 for plants from heavy shade, canopy light gaps and man-made clearings respectively. Light saturation of plants from clearings occurred at photosynthetic photon flux densities greater than 1000 mol m^–2 sec^–1 whereas plants from heavy shade environments became light saturated near 500 mol m^–2 sec^–1. Plants that normally occur in intermediate light environments were intermediate in light saturation levels. Mean maximum stomatal conductances ranged from 1.0 to 7.3 mm sec^–1 and followed a pattern similar to photosynthetic rates.     

The meiobenthos of five mangrove vegetation types in Gazi Bay,Kenya

The vertical distribution of meiofauna in the sediments ofAvicennia marina,Bruguiera gymnorrhiza,Ceriops tagal,Rhizophora mucronata andSonneratia alba at Gazi Bay (Kenya), is described. Seventeen taxa were observed, with highest densities in the sediments ofBruguiera (6707 ind. 10 cm^–2), followed byRhizophora (3998 ind. 10 cm^–2),Avicennia (3442 ind. 10 cm^–2),Sonneratia (2889 ind. 10 cm^–2) andCeriops (1976 ind. 10 cm^–2). Nematodes accounted for up to 95% of total densities; other common taxa were copepods, turbellarians, oligochaetes, polychaetes, ostracods and rotifers. High densities occurred to about 20 cm depth in the sediment. EspeciallyCeriops sediments show still high densities of nematodes (342 ind. 10 cm^–2) and copepods (11 ind. 10 cm^–2) in the deepest layer (15–22 cm). Particle size and oxygen conditions were major factors influencing meiobenthic distribution;Uca burrows had a major impact on distribution and abundance of meiofauna.     

Macroinvertebrate assemblages in Andean Patagonian rivers and streams: environmental relationships

Macroinvertebrate communities from 29 streams and rivers of the mountain and the Andean Patagonian Plateau were analyzed. Samples were collected from six river basins, which were part of four different biozones of the Patagonian Ecoregion. Samples from mountain streams were dominated by Plecoptera, Ephemeroptera, Trichoptera and Diptera, while plateau rivers where mainly Diptera, Oligochaeta and Mollusca. Total invertebrate abundance ranged from 7 to 12249 ind.m^–2. Elmidae, Paratrichocladius, Chironomus, Smicridea annulicornis, Parasericostoma ovale and Meridialaris laminata were the most abundant insect taxa, while Nais communis and Hyalella curvispina were the most abundant non-insect taxa. Species-environmental relationships were examined using Canonical Correspondence Analysis. Current speed, conductivity, substrate size and abundance of aquatic plants, were identified as the major variables structuring faunal assemblages. Regression analyses revealed that species richness was negatively correlated with latitude, and positively correlated with water temperature and altitude. Macroivertebrate abundance increased with conductivity, altitude and water temperature. These results suggest that habitat heterogeneity was the strongest predictor of macroinvertebrate assemblages, but species richness could be predicted at a landscape scale using topographical and climatic features.     

Compartment analysis of plant cells by means of turgor pressure relaxation: II. Experimental results onChara corallina

Summary The three-compartment model (paper I) described turgor pressure relaxations as a sum of two exponential functions. The predicted shape of the curves could be confirmed inChara corallina by improving the recording and processing of data. An evaluation on the basis of the three-compartment model provided values for the hydraulic conductivity of the plasmalemma ofLp _p=2×10^–5 to 4×10^–5 cm sec^–1 bar^–1 and ofLp _i=3×10^–5 to 1×10^–4 cm sec^–1 bar^–1 for the tonoplast (assuming the area to be 90% of the plasmalemma area). The mean proportion of the total volume occupied by the cytoplasm was calculated to be 9%. This value is consistent with the concept of a highly vacuolated cell. Other explanations for the biphasic relaxation curves are discussed.     

The reaction of larvae of Simulium noelleri (Diptera) to different current velocities

Larvae of Simulium noelleri Friederichs aggregate at high population densities (more than 10² cm^–2) on sluices, dams, and spillways. Experiments were conducted in a laboratory trough to assess the reaction of larvae to different current velocities (velocities ranged from 5–49 cm s^–1). In the lower part of the range of water velocities used, larvae moved a greater distance upstream from where they had been located. Larger larvae always showed a greater tendency to move than did smaller larvae, whatever the velocity. This intraspecific variation in reaction to different current velocities allows the aggregation of larvae of mixed sizes at suitable sites, smaller individuals being occluded by those that are larger.     

Living conditions, abundance and biomass of under-ice fauna in the Storfjord area (western Barents Sea, Arctic) in late winter (March 2003)

The under-ice habitat and fauna were studied during a typical winter situation at three stations in the western Barents Sea. Dense pack ice (7–10/10) prevailed and ice thickness ranged over <0.1–1.6 m covered by <0.1–0.6 m of snow. Air temperatures ranged between –1.8 and –27.5°C. The ice undersides were level, white and smooth. Temperature and salinity profiles in the under-ice water (0–5 m depth) were not stratified (T=–1.9 to –2.0°C and S=34.2–34.7). Concentrations of inorganic nutrients were high and concentrations of algal pigments were very low (0.02 g chlorophyll a l^–1), indicating the state of biological winter. Contents of particulate organic carbon and nitrogen ranged over 84.2–241.3 and 5.3–16.4 g l^–1, respectively, the C/N ratio over 11.2–15.5 pointing to the dominance of detritus in the under-ice water. Abundances of amphipods at the ice underside were lower than in other seasons: 0–1.8 ind. m^–2 for Apherusa glacialis, 0–0.7 ind. m^–2 for Onisimus spp., and 0–0.8 ind. m^–2 for Gammarus wilkitzkii. A total of 22 metazoan taxa were found in the under-ice water, with copepods as the most diverse and numerous group. Total abundances ranged over 181–2,487 ind. m^–3 (biomass: 70–2,439 g C m^–3), showing lower values than in spring, summer and autumn. The dominant species was the calanoid copepod Pseudocalanus minutus (34–1,485 ind. m^–3), contributing 19–65% to total abundances, followed by copepod nauplii (85–548 ind. m^–3) and the cyclopoid copepod Oithona similis (44–262 ind. m^–3). Sympagic (ice-associated) organisms occurred only rarely in the under-ice water layer.     

Assessing the current related heterogeneity and diversity patterns of benthic diatom communities in a turbid and a clear water river

Benthic diatoms were sampled in two rapids, in a turbid South-Finnish river (R. Keravanjoki, 22 FTU) and a clear water river in eastern Finland (R. Vaikkojoki, 4 FTU), to evaluate the diversity and spatial distribution patterns of diatom communities and especially their relationships to current velocity. In both rapids, epilithic diatoms were sampled in 15 sampling squares within three current velocity classes (10 cm s^–1, 40 cm s^–1 and 100 cm s^–1). The sampling squares had significantly different diatom communities in the three current velocity classes at both sampling sites, however, separation of the communities was much more pronounced in the clear water river (p < 0.001) than in the turbid river (p < 0.05). In the clear water river, communities at the highest velocity were highly different from those at the lower velocities. On the other hand, in the turbid river, communities were more similar at all velocities. Significant (p < 0.05) indicators for highest current velocity in the clear water R. Vaikkojoki were Fragilaria capucina var. gracilis Hustedt, F. capucina var. rumpens Lange-Bertalot and Meridion circulare Agardh. There were no significant indicators for high current velocity in the turbid R. Keravanjoki. Cocconeis placentula Ehr., Cymbella sinuata Gregory and Navicula lanceolata (Agardh) Ehr. were the three most abundant species in the highest velocity. This study showed that diatom community was highly specialized but low in diversity at the highest velocity in the clear water river. This pattern was not seen in the turbid R. Keravanjoki, indicating that the diatom community could withstand at the higher current velocities. In addition, the results show the importance of sampling in a variety of current regimes, particularly in clear water rivers, in order to properly assess the diatom diversity and community of a river section.     

Pulsed nuclear magnetic resonance study of39K within Halobacteria

Summary The³⁹K contents of isolated pellets and supernatant solutions from suspensions ofHalobacterium halobium were studied at 21–22°C by pulsed NMR spectroscopy. The rates of transverse relaxation were measured directly from the free induction decay (FID). The rate of longitudinal relaxation was measured by studying the FID after pairs of pulses of approximately 90°. Care was exercised to minimize the effect of magnetic field inhomogeneity; its contribution to the FID was approximately 25–30 sec^–1. The transverse relaxation process was found to consist of at least two components, whose rates were 321–449 sec^–1 and 1,122–2,067 sec^–1. In one preparation where the longitudinal relaxation process was studied, the data could be well fit to a single exponential relaxing at 253±33 (mean ±95% confidence limits) sec^–1. Comparison of the relative intensities of the NMR signals with the results of atomic absorption photometric analyses indicated that the great bulk of the intracellular³⁹K was detected by the NMR techniques used. The data obtained from the current NMR ofH. halobium are consistent with: (1) fractional binding of <3% of the total intracellular K⁺, (2) a small ordering factor characterizing all of the intracellular K⁺, or (3) some combination of the two.     

The summer zooplankton community at South Georgia: biomass,vertical migration and grazing

Zooplankton abundance and biomass were determined during January 1990 at two stations to the north-west of South Georgia using a Longhurst Hardy Plankton Recorder (LHPR). At both shelf and oceanic station sites, zooplankton biomass, (excluding Euphausia superba), was found to be ca. 13 g dry mass m^–2. Copepods and small euphausiids dominated the catches. These estimates are over 4 times higher than values generally reported for the Southern Ocean and may reflect firstly, the high productivity of the study area, secondly, the time of year, summer, when biomass for many species is maximal, and thirdly, the high sampling efficiency of the LHPR. Principal components analysis disclosed similarities and differences between adjacent depth strata in terms of abundance, biomass and species composition. At both stations most variability occurred in the mixed layer (0–60 m) and thermocline (60–120 m) with depth horizons below this being more homogeneous. Diel migrations were observed for most taxa with abundance increasing in the mixed layer at night. At the oceanic station, species and higher taxa belonging to the mesopelagic community were generally well spread throughout this domain and, with the exception of Pleuromamma robusta and Metridia curticauda, showed little evidence of migration. The grazing impact of the epipelagic community (copepods and small euphausiids) was estimated to remove 3–4% of the microbial standing stock day^–1 and a conservative 25% and 56% of daily primary production at the oceanic and shelf stations respectively.     

Macro- and megabenthic communities in the high Arctic Canada Basin: initial findings

We investigated the macro- and megabenthos of the Canada Basin, one of the biologically least known areas of the Arctic deep sea. Macro-infauna was collected with 11 box cores at six stations ranging from 640 to 3,250 m water depth. Total abundance ranged from 3 to 265 individuals 0.04 m^–2 (75–6,625 individuals m^–2) and decreased with increasing depth. Biomass ranged from 0.04 to 228 mg wet weight 0.04 m^–2 (1–5,700 mg wet weight m^–2) and followed the same trend. Polychaetes, crustaceans and bivalves dominated the faunal densities, biomass and species numbers. At Northwind Ridge (800 and 1,800 m), apparently historic (~5,000 years BP) fish otolith deposits were abundant. The invertebrate epi-megafauna was qualitatively analyzed from 9.2 h of video and 853 still images collected at four remotely operated vehicle stations. The epifauna was dominated by polychaetes, crustaceans, echinoderms, cnidarians and fish, with most suspension-feeders occurring in the Northwind Ridge area. A total of 90 benthic invertebrate species/taxa from four biogeographic affinities were identified, including at least three new species of Isopoda. The evident low abundances and biomass are in agreement with findings from the Eurasian Arctic deep sea.     

Distribution of dominant calanoid copepod species in the Greenland sea during late fall

Summary Between 6 November and 12 December 1988, vertical distributions of Calanus finmarchicus, C. hyperboreus, C. glacialis and Metridia longa were studied at three stations in the Arctic water of the Greenland Sea Gyre (GSG) and compared with two stations in the Atlantic water (AW) of the Westspitsbergen Current. Nine depth strata down to 3,000 m were sampled. C.finmarchicus was most abundant in AW, C. hyperboreus in GSG, M. longa showed no preference and C. glacialis was rare everywhere. Stage composition differed with species and water mass. Vertical distribution varied also with water mass in all species but C. hyperboreus, which was always centered between 1,000 and 1,500 m. The other species were concentrated in the upper 300 m in AW and between 1,000 and 1,500 m in GSG, although not all stages followed this trend. Ontogenetic vertical migration leads to significant dislocations of organic matter from the euphotic zone to great depth in the Greenland Sea. In egg production experiments, C. hyperboreus spawned up to 149 eggs female^–1 day^–1. Implications of stage composition, sex ratio, and gonad maturation on copepod life cycles are discussed.     

Abundance,population dynamics and production of Chironomidae (Diptera) in an ul traoligotrophic lake in South Greenland

Macrozoobenthos of the ultraoligotrophic Lake 95 (61°N, 46°W, 8 ha, z_max=18 m, ) is composed of about 14 taxa dominated by 12 Chironomidae species. Abundance, life cycle, biomass and production were estimated for the six dominant taxa. Abundance declined fromca. 4150 at 2.5 m depth toca. 1400 ind m^–2 at 16 m depth and averagedca. 3200 ind m^–2 on a lakewide basis. By numbers,Heterotrissocladius changi andH. oliveri dominated the average fauna.H. changi was common at the 2.5 m and 5 m depth stations, whereasH. oliveri dominated from 5 m depth downwards. Chironomids showed mainly a 1-yr life cycle, but apparently bothHeterotrissocladius species had two contemporary cohorts with emergence in midsummer and late autumn/early spring, respectively. Average annual ratio was 4.2 and 4.6 forH. oliveri andH. changi, respectively. Annual production varied from 0.3 g ash-free dry weight (AFDW) m^–2 y^–1 at 16 m depth to 1.6 g AFDW m^–2 y^–1 at 2.5 m depthH. changi contributed 45%, fiveMicropsectra spp. 17% andH. oliveri 15% to total average production, which on a lakewide basis wasca. 1.1 g AFDW or 25 kJ m^–2 y^–1. Lake 95 thus belongs at the very low end of measured lake zoobenthic productions, which range from 10 kJ m^–2 y^–1 in Arctic lakes toca. 1600 kJ m^–2 y^–1 in highly eutrophic shallow lakes.     

Use of artificial eggs in studies of washout depth and drift distance for salmonid eggs

Colour-coded artificial trout eggs were used in investigations of washout depth in a natural stream and of drift distance relative to water velocity in an experimental channel and in a section of natural stream.Washout depth was studied in a spawning riffle of a stream whose bankful discharge is 5.6 m³ s^–1. During an experiment when spates never exceeded 6.5 m³ s^–1 egg washout was severe at 5 cm depth within the gravel, variable at 10 cm and negligible at 15 cm. During another experiment when a spate of 9.0 m³ s^–1 (return period 10–20 years) occurred, washout was severe at 5 and 10 cm depth and variable at 15 cm. There was also evidence that some eggs were moved short distances downstream within the gravel without being washed out.Within experimental channels, drift distance could be predicted from multiple regressions relating logarithms of water velocity, percentage of eggs settled and distance from point of release. At a water velocity of 100 cm s^–1 at 0.6 depth, 50% of eggs would settle within 8 m of the point of release. At water velocities of 75 to 100 cm s^–1 drifting eggs would, on average, travel at c. 60% of water velocity and make 1 to 2 bed contacts m^–1 of travel.A similar multiple regression can be applied to data from a natural stream channel. It predicts much larger drift distances (50% settled in 42 m at 100 cm s ^–1 ). However, in the natural channel, settlement appears aggregated and the validity of the concept of permanent settlement is in doubt.     

An acoustic tracking of a megamouth shark, Megachasma pelagios: a crepuscular vertical migrator

A 4.9 m TL megamouth shark, only the sixth specimen known to science, was tracked continuously for 50.5 h, during which it exhibited distinct vertical migrations at the dawn and dusk transitions. The male shark was captured on 21 October 1990 in a drift gill net off Dana Point, California, restrained overnight in a harbor, and released at sea the next afternoon. Horizontally, the shark moved slowly southward, covering 62 km on a relatively straight path with no significant diel changes. For the major part of the tracking, its rate of movement was 1.15 km h^–1, as determined from positions at 15 min intervals. Considering a probable head current of 10–25 cm sec^–1, its estimated through-the-water swimming speed was more likely 1.5–2.1 km h^–1 (X¯ = 1.8, representing 0.1 body lengths sec^–1). Vertically, the shark stayed shallow at night (12–25 m depth range, X¯ = 17) and deep during the days (120–166 m, X¯ = 149) but still well above the bottom at 700–850 m. The four twilight depth-change events were very distinct and always spanned the times of sunset or sunrise. The ascent and descent profiles are a reasonble match to isolumes on the order of 0.4 lux for an extinction coefficient (0.07) calculated from water transparency measurements. Furthermore, the steepest parts of the shark's profiles correspond closely to the times of maximum rate-of-change of illumination. These findings suggest that, except during nights, the shark's chosen depth was to a large degree determined by light level.