Mark C. Benfield

From the Hensen net toward four-dimensional biological oceanography

Abstract The development of quantitative zooplankton collecting systems began with Hensen, 1887 , Hensen, 1895 ). Non-opening closing nets, opening closing nets (mostly messenger based), high-speed samplers, and planktobenthos net systems all had their start in his era — the late 1800s and early 1900s. This was also an era in which many of the fundamental questions about the structure and dynamics of the plankton in the worlds oceans were first posed. Fewer new systems were introduced between 1912 and 1950 apparently due in part to the two World Wars. The continuous plankton recorder stands out as a truly innovative device developed during this period ( Hardy 1926b Nature, London 118, 630 ). Resurgence in development of mechanically-based instruments occurred during the 1950s and 1960s. A new lineage of high-speed samplers, the Gulf series, began in the 1950s and a number of variants were developed in the 1960s and 1970s. Net systems specifically designed to collect neuston first appeared in the late 1950s. During the 1960s, many focused field and experimental tank experiments were carried out to investigate the hydrodynamics of nets, and much of our knowledge concerning net design and construction criteria was developed. The advent of reliable electrical conducting cables and electrically-based control systems during this same period gave rise first to a variety of cod-end samplers and then to the precursors of the acoustically and electronically-controlled multi-net systems and environmental sensors, which appeared in the 1970s. The decade of the 1970s saw a succession of multi-net systems based both on the Be multiple plankton sampler and on the Tucker trawl. The advent of the micro-computer stimulated and enabled the development of sophisticated control and data logging electronics for these systems in the 1980s. In the 1990s, acoustic and optical technologies gave rise to sensor systems that either complement multiple net systems or are deployed without nets. Multi-sensor systems with high data telemetry rates through electro-optical cable are now being deployed in towed bodies and on remotely operated vehicles. In the offing are new molecular technologies to identify species in situ, and realtime data analysis, image processing, and 3D/4D display. In the near future, it is likely that the use of multi-sensor systems deployed on autonomous vehicles will yield world wide coverage of the distribution and abundance of zooplankton.

Time to automate identification

Taxonomists should work with specialists in pattern recognition, machine learning and artificial intelligence, say Norman MacLeod, Mark Benfield and Phil Culverhouse — more accuracy and less drudgery will result.

Stable isotope indicators of movement and residency for brown shrimp (Farfantepenaeus aztecus) in coastal Louisiana marshscapes

Brown shrimp (Farfantepenaeus aztecus) are an important commercial aquatic species experiencing loss of inshore marsh nursery habitat in coastal Louisiana. To study inshore brown shrimp movements and identify aspects of essential habitat important for sustaining brown shrimp populations, we collected juvenile brown shrimp in April and May 2000, the time of annual maximum brown shrimp abundance, in a small 1-km2 marsh area on the central Louisiana coast. Drop sampling showed average shrimp densities of 1.6–2.4 m−2 in shallow marsh ponds and seining indicated lower densities of 0.5–0.9 m−2 in nearby shallow channel and open bay sites. Smaller shrimp (< 50 mm) fed disproportionately on benthic diatoms and small harpacticoid copepods, while large shrimp fed more frequently on larger-bodied amphipods and tanaids. We developed novel chemical approaches to estimate patterns of shrimp residency and movement using carbon and nitrogen stable isotopic determinations. Resident shrimp had isotopic values similar to average foods and showed consistent isotopic spacings between fast and slow turnover tissues. Residency was highest (47–55%) in ponds and shallow channel habitats and much less in open bays and deep channels (4–27%). There was sparse evidence for dietary specialization among individull shrimp. The results support the view that small 10–20 mm postlarval and juvenile brown shrimp arriving in estuaries from offshore waters continue movement through sub-optimal habitats (deep channels and open bays), but exhibit much less movement once an optimal habitat (marsh ponds or shallow channel margins) is reached. This study also indicated that combining estimates of shrimp densities, residency, growth rate, and mortality allows evaluation of the importance of different habitat types for shrimp production. Shallow ponds that in many ways resemble fertile aquaculture ponds appear to be hot spots for brown shrimp production, and coastal preservation and restoration efforts should focus on these areas as important for sustaining shrimp fisheries.

Determining dominant scatterers of sound in mixed zooplankton populations

High-frequency acoustic scattering techniques have been used to investigate dominant scatterers in mixed zooplankton populations. Volume backscattering was measured in the Gulf of Maine at 43, 120, 200, and 420 kHz. Zooplankton composition and size were determined using net and video sampling techniques, and water properties were determined using conductivity, temperature, and depth sensors. Dominant scatterers have been identified using recently developed scattering models for zooplankton and microstructure. Microstructure generally did not contribute to the scattering. At certain locations, gas-bearing zooplankton, that account for a small fraction of the total abundance and biomass, dominated the scattering at all frequencies. At these locations, acoustically inferred size agreed well with size determined from the net samples. Significant differences between the acoustic, net, and video estimates of abundance for these zooplankton are most likely due to limitations of the net and video techniques. No other type of biological scatterer ever dominated the scattering at all frequencies. Copepods, fluid-like zooplankton that account for most of the abundance and biomass, dominated at select locations only at the highest frequencies. At these locations, acoustically inferred abundance agreed well with net and video estimates. A general approach for the difficult problem of interpreting high-frequency acoustic scattering in mixed zooplankton populations is described.

High-frequency acoustic volume backscattering in the Georges Bank coastal region and its interpretation using scattering models

High-frequency (120 and 420 kHz) sound was used to survey sound scatterers in the water over Georges Bank. In addition to the biological sound scatterers (the plankton and micronekton), scattering associated with internal waves and suspended sediment was observed. Volume backscattering was more homogeneous in the vertical dimension (with occasional patches) in the shallow central portion of the Bank where there is significant mixing. In the deeper outer portion of the Bank where the water is stratified, volume backscattering was layered and internal waves modulated the vertical position of the layers in the pycnocline. The internal waves typically had amplitudes of 5-20 m, but sometimes much higher. Species composition and size data from samples of the animals and suspended sediment used in conjunction with acoustic scattering models revealed that throughout the region the animals generally dominate the scattering, but there are times and places where sand particles (suspended as high as up to the sea surface) can dominate. The source of the scattering in the internal waves is probably due to a combination of both animals and sound-speed microstructure. Determination of their relative contributions requires further study.

Sizing ocean giants: patterns of intraspecific size variation in marine megafauna

What are the greatest sizes that the largest marine megafauna obtain? This is a simple question with a difficult and complex answer. Many of the largest-sized species occur in the world’s oceans. For many of these, rarity, remoteness, and quite simply the logistics of measuring these giants has made obtaining accurate size measurements difficult. Inaccurate reports of maximum sizes run rampant through the scientific literature and popular media. Moreover, how intraspecific variation in the body sizes of these animals relates to sex, population structure, the environment, and interactions with humans remains underappreciated. Here, we review and analyze body size for 25 ocean giants ranging across the animal kingdom. For each taxon we document body size for the largest known marine species of several clades. We also analyze intraspecific variation and identify the largest known individuals for each species. Where data allows, we analyze spatial and temporal intraspecific size variation. We also provide allometric scaling equations between different size measurements as resources to other researchers. In some cases, the lack of data prevents us from fully examining these topics and instead we specifically highlight these deficiencies and the barriers that exist for data collection. Overall, we found considerable variability in intraspecific size distributions from strongly left- to strongly right-skewed. We provide several allometric equations that allow for estimation of total lengths and weights from more easily obtained measurements. In several cases, we also quantify considerable geographic variation and decreases in size likely attributed to humans.

Comparison of multifrequency acoustic and in situ measurements of zooplankton abundances in Knight Inlet, British Columbia

An investigation of midwater zooplankton aggregations in a coastal fjord was conducted in November 2002. This study focused on quantitative comparisons between a calibrated, three-frequency (38, 120, and 200 kHz) vessel-based echo-sounder, a multinet towed zooplankton sampler (BIONESS), and a high-resolution underwater camera (ZOOVIS). Daytime layers of euphausiids and amphipods near 70-90-m depth were observed in lower parts of the inlet, especially concentrated by tidal flows around a sill. Quantitative backscatter measurements of euphausiids and amphipods, combined with in situ size and abundance estimates, and using an assumed tilt-angle distribution, were in agreement with averaged fluid-cylinder scattering models produced by Stanton and Chu [ICES J. Mar. Sci. 57, 793-807, (2000)]. Acoustic measurements of physonect siphonophores in the upper inlet were found to have a strong 38-kHz scattering strength, in agreement with a damped bubble scattering model using a diameter of 0.4 mm. In relatively dense euphausiid layers, ZOOVIS abundance estimates were found to be a factor of 2 to 4 higher than the acoustic estimates, potentially due to deviations from assumed euphausiid orientation. Nocturnal near-surface euphausiid scattering exhibited a strong (15 dB) and rapid (seconds) sensitivity to vessel lights, interpreted as due to changing animal orientation.

Three-Dimensional Acoustic Visualization of Zooplankton Patchiness

Abstract Acoustic data were collected and visualized to characterize the 3-dimensional patchiness of zooplankton at a thermally stratified site on Georges Bank. The work was carried out as part of a field study conducted to examine the effects of springtime water-column stratification on the distributions of zooplankton and larval fish on the Bank. The acoustic data were acquired as the ship steamed a survey grid relative to the track of a surface drifter with a subsurface drogue. Although quite irregular in geographical coordinate space, the ship’s track relative to the moving water closely matched the intended grid pattern once the drifter’s movement in the tidal flow was taken into account. After changing coordinate systems to compensate for tidal advection, the acoustic data set was transformed from its curtain-like distribution in 3-dimensional space to a volumetric distribution. Two-dimensional point kriging was performed on the irregularly spaced data from each 2-m-thick depth stratum to produce a series of 2-dimensional, regularly spaced data grids. These data grids were then stacked to construct the 3-dimensional data grid required for volumetric visualization. A similar procedure was followed with the error variance values produced at each grid point through kriging to construct a 3-dimensional, volumetric distribution of the error variance. To examine zooplankton patchiness within the surveyed volume of water, isosurfaces corresponding to specific levels of acoustic backscatter were highlighted in the visualization. The 3-dimensional distribution of error variance was used to control the opacity of the isosurfaces to provide an objective, visual approach for displaying the statistical confidence one can have in the patches detected. In this survey, the ship steamed directly over a large, southwest- to northeast-oriented patch of zooplankton on at least three different passes. It also steamed over several smaller patches. The vertically compressed nature of the patches and their high degree of spatial heterogeneity in the horizontal plane are characteristic of the zooplankton distributions found in the deeper, seasonally stratified portions of Georges Bank.

An empirical assessment of the consistency of taxonomic identifications

Abstract Plankton counting and analysis is essential in ecological study, yet scant literature exists as to the reliability of those counts and the consistency of the experts who make the counts. To assess how variable expert taxonomic identifications are, a set of six archived mesozooplankton samples from a series of Longhurst Hardy Plankton Recorder net hauls were counted by expert zooplankton analysts located at six marine laboratories. Sample identifications were repeated on two separate days with over 700 target specimens counted and identified on each day across the samples. Twenty percent of the analysts returned counts that varied by more than 10%. Thirty-three percent of analysts exhibited low identification consistencies, returning Intraclass Correlation Coefficient scores of less than 0.80. Statistical analyses of these data suggest that over 83% of the observed categorical count variance can be attributed to inconsistencies within analysts. We suggest this is the root cause of variation in expert specimen labelling consistency.

In Situ Video Observations of Two Manefishes (Perciformes: Caristiidae) in the Mesopelagic Zone of the Northern Gulf of Mexico

Abstract This paper describes direct video observations of two manefishes, likely Paracaristius sp., from the mesopelagic waters of the north-central Gulf of Mexico. One fish was observed with a remotely operated vehicle at a depth of 829 m by an industrial ROV as part of the SERPENT Project. The second was observed at 496 m from a manned submersible. Little is known about the behavior of manefishes because most records result from net-collected material. Our observation demonstrates that manefishes are capable of precise locomotory and posture control using extended, erect fins and that the pelvic fins of these fishes are extended in a parachute-like manner. Moreover, one of the specimens exhibited an unusual vertical, sinusoidal oscillation of its caudal fin. One of the observations took place in association with a physonect siphonophore. These observations may include the deepest published record for a manefish in the Gulf of Mexico.