Mesopelagic fish collected during Maria S. Merian cruise MSM26, spring 2013

Mesopelagic fish were collected using a 1 m**2 Double-MOCNESS (Multiple Opening and Closing Net and Environmental Sensing System) and 4.5 m**2 IKMT (Isaacs-Kidd midwater trawl). The main portion of the IKMT was 20 mm knotted nylon, and the tail bag was 3 mm knotless nylon. Oblique IKMT tows were made to a maximum depth of 500 m at a tow speed of 3.5 knots. The original cruise plan intended for nighttime IKMT tows, but tow times varied due to operational constraints. The MOCNESS was equipped with 20 nets of 333 µm mesh size; 10 nets per side. The towing speed was 2 knots. Samples were collected to a maximum depth of 1250 m. The first oblique nets sampled from the surface to the max depth, and the other nets sampled depth stratified bins of the water column. MOCNESS hauls were performed during day and night to investigate diel vertical migrations. Mesoplelagic fish were processed on board. All fish were picked from all IKMT nets, most oblique MOCNESS nets, and the left side nets of the depth stratified MOCNESS samples. The Depth stratified nets from the right side of the MOCNESS frame were preserved in 5 % formalin for future quantitative analyses of the nekton. Fish were identified to the lowest possible taxa using Whitehead et al. (1984) and Fahay (2007). Standard length of each fish was measured to the nearest 0.1 mm using a digital caliper. Measured and identified fish were frozen in an -80 °C freezer, and shipped to the University of Hamburg at the end of the cruise.

Abundance mesopelagic fish collected during Maria S. Merian cruise MSM26, spring 2013

Mesopelagic fish were collected using a 1 m**2 Double-MOCNESS (Multiple Opening and Closing Net and Environmental Sensing System) and 4.5 m**2 IKMT (Isaacs-Kidd midwater trawl). The main portion of the IKMT was 20 mm knotted nylon, and the tail bag was 3 mm knotless nylon. Oblique IKMT tows were made to a maximum depth of 500 m at a tow speed of 3.5 knots. The original cruise plan intended for nighttime IKMT tows, but tow times varied due to operational constraints. The MOCNESS was equipped with 20 nets of 333 µm mesh size; 10 nets per side. The towing speed was 2 knots. Samples were collected to a maximum depth of 1250 m. The first oblique nets sampled from the surface to the max depth, and the other nets sampled depth stratified bins of the water column. MOCNESS hauls were performed during day and night to investigate diel vertical migrations. Mesoplelagic fish were processed on board. All fish were picked from all IKMT nets, most oblique MOCNESS nets, and the left side nets of the depth stratified MOCNESS samples. The Depth stratified nets from the right side of the MOCNESS frame were preserved in 5 % formalin for future quantitative analyses of the nekton. Fish were identified to the lowest possible taxa using Whitehead et al. (1984) and Fahay (2007). Standard length of each fish was measured to the nearest 0.1 mm using a digital caliper. Measured and identified fish were frozen in an -80 °C freezer, and shipped to the University of Hamburg at the end of the cruise.

Micronekton of the North Pacific

1. INTRODUCTION 1.1 Working Group History 2. SPECIES COMPOSITION AND DISTRIBUTION PATTERNS RELATED TO WATER MASSES 2.1 Mesopelagic Fishes 2.1.1 Dominant families 2.1.2 Large-scale feeding and/or spawning migration or expatriation? 2.1.3 Definition of water masses 2.1.4 Species composition 2.2 Crustacean Micronekton 2.2.1 Euphausiids 2.2.2 Mysids and decapods 2.3 Cephalopod Micronekton 2.3.1 Family Enoploteuthidae 2.3.2 Family Gonatidae 2.3.3 Family Onychoteuthidae 2.3.4 Family Pyroteuthidae 2.3.5 Other cephalopods 3. VERTICAL DISTRIBUTION PATTERNS 3.1 Mesopelagic Fishes 3.1.1 Significance of diel vertical migration 3.1.2 DVM patterns 3.1.3 Ontogenetic change in DVM patterns 3.2 Crustacean Micronekton 3.3 Cephalopod Micronekton 4. BIOMASS PATTERNS 4.1 Micronektonic Fish 5. LIFE HISTORY 5.1 Fish Micronekton 5.1.1 Age and growth 5.1.2 Production 5.1.3 Reproduction 5.1.4 Mortality 5.2 Crustacean Micronekton 5.2.1 Age and growth 5.2.2 Production 5.2.3 Reproduction and early life history 5.2.4 Mortality 5.3 Cephalopod Micronekton 5.3.1 Age and growth 5.3.2 Production 5.3.3 Reproduction and early life history 5.3.4 Mortality 6. ECOLOGICAL RELATIONS 6.1 Feeding Habits 6.1.1 Fish micronekton 6.1.2 Crustacean micronekton 6.1.3 Cephalopod micronekton 6.2 Estimating the Impact of Micronekton Predation on Zooplankton 6.2.1 Predation by micronektonic fish 6.3 Predators 6.3.1 Cephalopods 6.3.2 Elasmobranchs 6.3.3 Osteichthyes 6.3.4 Seabirds 6.3.5 Pinnipeds 6.3.6 Cetaceans 6.3.7 Human consumption 6.4 Predation Rate 6.5 Ecosystem Perspectives 6.6 Interactions between Micronekton and Shallow Topographies 7. SAMPLING CONSIDERATIONS 7.1 Net Trawling 7.1.1 Sampling gears 7.1.2 Sampling of surface migratory myctophids 7.1.3 Commercial-sized trawl sampling 7.1.4 Sampling of euphausiids and pelagic decapods 7.2 Acoustic Sampling 7.2.1 Acoustic theory and usage 7.3 Video Observations (Submersible and ROV) 8. SUMMARY OF PRESENT STATE OF KNOWLEDGE 8.1 Fish Micronekton 8.2 Crustacean Micronekton 8.3 Cephalopod Micronekton 9. RECOMMENDATIONS 10. REFERENCES 11. APPENDICES (122 page document)

Abundance and Distribution of Ichthyoplankton along an Inshore-Offshore Transect in Onslow Bay, North Carolina

The distribution and abundance of ichthyoplankton was investigated from November 1979 to March 1980 along a transect from coastal to continental slope waters in Onslow Bay, North Carolina. Representatives of 66 families were collected; 24 of which were tropical families, a category that also includes families of typically oceanic and deep-sea fishes. Larvae of tropical species were collected in coastal and shelf waters, demonstrating the intrusion of Gulf Stream waters onto the continental shelf. From December through March, frontal waters that separated cold open-shelf surface waters from warm Gulf Stream surface waters were observed. Higher abundances of fish larvae were sometimes, but not consistently, associated with frontal waters. A great diversity of taxa was collected in offshore waters, and densities of larvae were low in coastal waters; low densities were attributed to gear selectivity rather than low larval abundance. Larvae of commercially and recreationally important estuarine-dependent species, especially Leiostomus xanthus and Micropogonias undulatus, were dominant components of the ichthyoplankton. Representatives of the families Bothidae, Clupeidae, Gadidae, Gonostomatidae, Myctophidae, Ophidiidae, and Sparidae were also important components of the ichthyoplankton. Larvae of species representing two strikingly different life history types-mesopelagic and estuarine-dependent frequently cooccurred.(PDF file contains 32 pages.)

Postrelease survival, vertical and horizontal movements, and thermal habitats of five species of pelagic sharks in the central Pacific Ocean

From 2001 to 2006, 71 pop-up satellite archival tags (PSATs) were deployed on five species of pelagic shark (blue shark [Prionace glauca]; shortfin mako [Isurus oxyrinchus]; silky shark [Carcharhinus falciformis]; oceanic whitetip shark [C. longimanus]; and bigeye thresher [Alopias superciliosus]) in the central Pacific Ocean to determine species-specific movement patterns and survival rates after release from longline fishing gear. Only a single postrelease mortality could be unequivocally documented: a male blue shark which succumbed seven days after release. Meta-analysis of published reports and the current study (n=78 reporting PSATs) indicated that the summary effect of postrelease mortality for blue sharks was 15% (95% CI, 8.5–25.1%) and suggested that catch-and-release in longline fisheries can be a viable management tool to protect parental biomass in shark populations. Pelagic sharks displayed species-specific depth and temperature ranges, although with significant individual temporal and spatial variability in vertical movement patterns, which were also punctuated by stochastic events (e.g., El Niño-Southern Oscillation). Pelagic species can be separated into three broad groups based on daytime temperature preferences by using the unweighted pair-group method with arithmetic averaging clustering on a Kolmogorov-Smirnov Dmax distance matrix: 1) epipelagic species (silky and oceanic whitetip sharks), which spent >95% of their time at temperatures within 2°C of sea surface temperature; 2) mesopelagic-I species (blue sharks and shortfin makos, which spent 95% of their time at temperatures from 9.7° to 26.9°C and from 9.4° to 25.0°C, respectively; and 3) mesopelagic-II species (bigeye threshers), which spent 95% of their time at temperatures from 6.7° to 21.2°C. Distinct thermal niche partitioning based on body size and latitude was also evident within epipelagic species.

Spatial and temporal variability in the distribution and abundance of larval and juvenile fishes associated with pelagic Sargassum

A survey of the larval and juvenile fishes associated with the pelagic Sargassum habitat in the South Atlantic Bight and adjacent western Atlantic Ocean was conducted from July 1991 through March 1993. Fishes representing 104 taxonomic categories were identified, including reef fishes, coastal demersal, coastal pelagic, epipelagic and mesopelagic species. The most important families were Balistidae and Carangidae, each represented by 15 species. Species composition, species diversity and abundance varied both seasonally and regionally. Diversity was highest during spring through fall over the outer continental shelf and in the Gulf Stream. Abundance decreased from spring through winter and from the continental shelf into offshore waters. The numbers of fishes and fish biomass were found to be positively correlated with the wet weight of algae in most cases examined. The results of this study will be useful to fisheries managers assessing the potential impacts of commercial Sargassum harvesting in the region.

Food habits and consumption rates of common dolphinfish (Coryphaena hippurus) in the eastern Pacific Ocean

An ecosystem approach to fisheries management requires an understanding of the impact of predatory fishes on the underlying prey resources. Defining trophic connections and measuring rates of food consumption by apex predators lays the groundwork for gaining insight into the role of predators and commercial fisheries in influencing food web structure and ecosystem dynamics.We analyzed the stomach contents of 545 common dolphinfish (Coryphaena hippurus) sampled from 74 sets of tuna purse-seine vessels fishing in the eastern Pacific Ocean (EPO) over a 22-month period. Stomach fullness of these dolphinfish and digestion state of the prey indicated that diel feeding periodicity varied by area and may be related to the digestibility and energy content of the prey. Common dolphinfish in the EPO appear to feed at night, as well as during the daytime. We analyzed prey importance by weight, numbers, and frequency of occurrence for five regions of the EPO. Prey importance varied by area. Flyingfishes, epipelagic cephalopods, tetraodontiform fishes, several mesopelagic fishes, Auxis spp., and gempylid fishes predominated in the diet. Ratios of prey length to predator length ranged from 0.014 to 0.720. Consumption-rate estimates averaged 5.6% of body weight per day. Stratified by sex, area, and length class, daily rations ranged up to 9.6% for large males and up to 19.8% for small dolphinfish in the east area (0–15°N, 111°W–coastline). Because common dolphinfish exert substantial predation pressure on several important prey groups, we concluded that their feeding ecology provides important clues to the pelagic food web and ecosystem structure in the EPO.

Trophic role of small cetaceans and seals in Irish waters

Understanding the role of marine mammals in specific ecosystems and their interactions with fisheries involves, inter alia, an understanding of their diet and dietary requirements. In this thesis, the foraging ecology of seven marine mammal species that regularly occur in Irish waters was investigated by reconstructing diet using hard parts from digestive tracts and scats. Of the species examined, two (striped and Atlantic white-sided dolphin) can be considered offshore species or species inhabiting neritic waters, while five others usually inhabit more coastal areas (white-beaked dolphin, harbour porpoise, harbour seal and grey seal); the last species studied was the bottlenose dolphin whose population structure is more complex, with coastal and offshore populations. A total of 13,028 prey items from at least 81 different species (62 fish species, 14 cephalopods, four crustaceans, and a tunicate) were identified. 28% of the fish species were identified using bones other than otoliths, highlighting the importance of using all identifiable structures to reconstruct diet. Individually, each species of marine mammal presented a high diversity of prey taxa, but the locally abundant Trisopterus spp. were found to be the most important prey item for all species, indicating that Trisopterus spp. is probably a key species in understanding the role of these predators in Irish waters. In the coastal marine mammals, other Gadiformes species (haddock, pollack, saithe, whiting) also contributed substantially to the diet; in contrast, in pelagic or less coastal marine mammals, prey was largely comprised of planktivorous fish, such as Atlantic mackerel, horse mackerel, blue whiting, and mesopelagic prey. Striped dolphins and Atlantic white-sided dolphins are offshore small cetaceans foraging in neritic waters. Differences between the diet of striped dolphins collected in drift nets targeting tuna and stranded on Irish coasts showed a complex foraging behaviour; the diet information shows that although this dolphin forages mainly in oceanic waters it may occasionally forage on the continental shelf, feeding on available prey. The Atlantic white-sided dolphin diet showed that this species prefers to feed over the continental edge, where planktivorous fish are abundant. Some resource partitioning was found in bottlenose dolphins in Irish waters consistent with previous genetic and stable isotope analysis studies. Bottlenose dolphins in Irish waters appears to be generalist feeders consuming more than 30 prey species, however most of the diet comprised a few locally abundant species, especially gadoid fish including haddock/pollack/saithe group and Trisopterus spp., but the contribution of Atlantic hake, conger eels and the pelagic planktivorous horse mackerel were also important. Stomach content information suggests that three different feeding behaviours might occur in bottlenose dolphin populations in Irish waters; firstly a coastal behaviour, with animals feeding on prey that mainly inhabit areas close to the coast; secondly an offshore behaviour where dolphins feed on offshore species such as squid or mesopelagic fish; and a third more complex behaviour that involves movements over the continental shelf and close to the shelf edge. The other three coastal marine mammal species (harbour porpoise, harbour seal and grey seal) were found to be feeding on similar prey and competition for food resources among these sympatric species might occur. Both species of seals were found to have a high overlap (more than 80%) in their diet composition, but while grey seals feed on large fish (>110mm), harbour seals feed mostly on smaller fish (<110mm), suggesting some spatial segregation in foraging. Harbour porpoises and grey seals are potentially competing for the same food resource but some differences in prey species were found and some habitat partitioning might occur. Direct interaction (by catch) between dolphins and fisheries was detected in all species. Most of the prey found in the stomach contents from both stranded and by catch dolphins were smaller sizes than those targeted by commercial fisheries. In fact, the total annual food consumption of the species studied was found to be very small (225,160 tonnes) in comparison to fishery landings for the same area (~2 million tonnes). However, marine mammal species might be indirectly interacting with fisheries, removing forage fish. Incorporating the dietary information obtained from the four coastal species, an ECOPATH food web model was established for the Irish Sea, based on data from 2004. Five trophic levels were found, with bottlenose dolphins and grey and harbour seals occurring at the highest trophic level. A comparison with a previous model based on 1973 data suggests that while the overall Irish Sea ecosystem appears to be “maturing”, some indices indicate that the 2004 fishery was less efficient and was targeting fish at higher trophic levels than in 1973, which is reflected in the mean trophic level of the catch. Depletion or substantial decrease of some of the Irish Sea fish stocks has resulted in a significant decline in landings in this area. The integration of diet information in mass-balance models to construct ecosystem food-webs will help to understand the trophic role of these apex predators within the ecosystem.

Seasonal movements and behaviour of basking sharks from archival tagging: No evidence of winter hibernation

Habitat selection processes in highly migratory animals such as sharks and whales are important to understand because they influence patterns of distribution, availability and therefore catch rates. However, spatial strategies remain poorly understood over seasonal scales in most species, including, most notably, the plankton-feeding basking shark Cetorhinus maximus. It was proposed nearly 50 yr ago that this globally distributed species migrates from coastal summer-feeding areas of the northeast Atlantic to hibernate during winter in deep water on the bottom of continental-shelf slopes. This view has perpetuated in the literature even though the 'hibernation theory' has not been tested directly. We have now tracked basking sharks for the first time over seasonal scales (1.7 to 6.5 mo) using 'pop-up' satellite archival transmitters. We show that they do not hibernate during winter but instead undertake extensive horizontal (up to 3400 km) and vertical (> 750 m depth) movements to utilise productive continental-shelf and shelf-edge habitats during summer, autumn and winter. They travel long distances (390 to 460 km) to locate temporally discrete productivity 'hotspots' at shelf-break fronts, but at no time were prolonged movements into open-ocean regions away from shelf waters observed. Basking sharks have a very broad vertical diving range and can dive beyond the known range of planktivorous whales. Our study suggests this species can exploit shelf and slope-associated zooplankton communities in mesopelagic (200 to 1000 m) as well as epipelagic habitat (0 to 200 m).

DISCOVERY 2010: Spatial and temporal variability in a dynamic polar ecosystem

The Scotia Sea has been a focus of biological- and physical oceanographic study since the Discovery expeditions in the early 1900s. It is a physically energetic region with some of the highest levels of productivity in the Southern Ocean. It is also a region within which there have been greater than average levels of change in upper water column temperature. We describe the results of three cruises transecting the central Scotia Sea from south to north in consecutive years and covering spring, summer and autumn periods. We also report on some community level syntheses using both current-day and historical data from this region. A wide range of parameters were measured during the field campaigns, covering the physical oceanography of the region, air–sea CO2 fluxes, macro- and micronutrient concentrations, the composition and biomass of the nano-, micro- and mesoplankton communities, and the distribution and biomass of Antarctic krill and mesopelagic fish. Process studies examined the effect of iron-stress on the physiology of primary producers, reproduction and egestion in Antarctic krill and the transfer of stable isotopes between trophic layers, from primary consumers up to birds and seals. Community level syntheses included an examination of the biomass-spectra, food-web modelling, spatial analysis of multiple trophic layers and historical species distributions. The spatial analyses in particular identified two distinct community types: a northern warmer water community and a southern cold community, their boundary being broadly consistent with the position of the Southern Antarctic Circumpolar Current Front (SACCF). Temperature and ice cover appeared to be the dominant, over-riding factors in driving this pattern. Extensive phytoplankton blooms were a major feature of the surveys, and were persistent in areas such as South Georgia. In situ and bioassay measurements emphasised the important role of iron inputs as facilitators of these blooms. Based on seasonal DIC deficits, the South Georgia bloom was found to contain the strongest seasonal carbon uptake in the ice-free zone of the Southern Ocean. The surveys also encountered low-production, iron-limited regions, a situation more typical of the wider Southern Ocean. The response of primary and secondary consumers to spatial and temporal heterogeneity in production was complex. Many of the life-cycles of small pelagic organisms showed a close coupling to the seasonal cycle of food availability. For instance, Antarctic krill showed a dependence on early, non-ice-associated blooms to facilitate early reproduction. Strategies to buffer against environmental variability were also examined, such as the prevalence of multiyear life-cycles and variability in energy storage levels. Such traits were seen to influence the way in which Scotia Sea communities were structured, with biomass levels in the larger size classes being higher than in other ocean regions. Seasonal development also altered trophic function, with the trophic level of higher predators increasing through the course of the year as additional predator-prey interactions emerged in the lower trophic levels. Finally, our studies re-emphasised the role that the simple phytoplankton-krill-higher predator food chain plays in this Southern Ocean region, particularly south of the SACCF. To the north, alternative food chains, such as those involving copepods, macrozooplankton and mesopelagic fish, were increasingly important. Continued ocean warming in this region is likely to increase the prevalence of such alternative such food chains with Antarctic krill predicted to move southwards.

A blooming jellyfish in the northeast Atlantic and Mediterranean

A long-term time series of plankton records collected by the continuous plankton recorder (CPR) Survey in the northeast Atlantic indicates an increased occurrence of Cnidaria since 2002. In the years 2007 and 2008, outbreaks of the warm-temperate scyphomedusa, Pelagia noctiluca, appeared in CPR samples between 45° N to 58° N and 1° W to 26° W. Knowing the biology of this species and its occurrence in the adjacent Mediterranean Sea, we suggest that P. noctiluca may be exploiting recent hydroclimatic changes in the northeast Atlantic to increase its extent and intensity of outbreaks. In pelagic ecosystems, Cnidaria can affect fish recruitment negatively. Since P. noctiluca is a highly venomous species, outbreaks can also be detrimental to aquaculture and make bathing waters unusable, thus having profound ecological and socio-economic consequences.

Sedimentation of acantharian cysts in the Iceland Basin: Strontium as a ballast for deep ocean particle flux, and implications for acantharian reproductive strategies

Acantharian cysts were discovered in sediment trap samples from spring 2007 at 2000 m in the Iceland Basin. Although these single-celled organisms contribute to particulate organic matter flux in the upper mesopelagic, their contribution to bathypelagic particle flux has previously been found negligible. Four time-series sediment traps were deployed and all collected acantharian cysts, which are reproductive structures. Across all traps, cysts contributed on average 3-22%, and 4―24% of particulate organic carbon and nitrogen (POC and PON) flux, respectively, during three separate collection intervals (the maximum contribution in any one trap was 48% for POC and 59% for PON). Strontium (Sr) flux during these 6 weeks reached 3 mg m―2 d―1. The acantharian celestite (SrSO4) skeleton clearly does not always dissolve in the mesopelagic as often thought, and their cysts can contribute significantly to particle flux at bathypelagic depths during specific flux events. Their large size (∼ I mm) and mineral ballast result in a sinking rate of ∼ 500 m d―1; hence, they reach the bathypelagic before dissolving. Our findings are consistent with a vertical profile of salinity-normalized Sr concentration in the Iceland Basin, which shows a maximum at 1700 m. Profiles of salinity-normalized Sr concentration in the subarctic Pacific reach maxima at ≤ 1500 m, suggesting that Acantharia might contribute to the bathypelagic particle flux there as well. We hypothesize that Acantharia at high latitudes use rapid, deep sedimentation of reproductive cysts during phytoplankton blooms so that juveniles can exploit the large quantity of organic matter that sinks rapidly to the deep sea following a bloom.

The effect of the North Atlantic Subpolar Front as a boundary in pelagic biogeography decreases with increasing depth and organism size

Broad-scale patterns in the distribution of deep-sea pelagic species and communities are poorly known. An important question is whether biogeographic boundaries identified from surface features are important in the deep mesopelagic and bathypelagic. We present community analyses of discrete-depth samples of mesozooplankton and micronekton to full-ocean depth collected in the area where the Mid-Atlantic Ridge is crossed by the Subpolar Front. The results show that the distributional discontinuity associated with the front, which is strong near the surface, decreases with increasing depth. Both the frontal separation near the surface and the community convergence at increasing depths were clearer for mesozooplankton than for micronekton.

The effect of the North Atlantic Subpolar Front as a boundary in pelagic biogeography decreases with increasing depth and organism size

Broad-scale patterns in the distribution of deep-sea pelagic species and communities are poorly known. An important question is whether biogeographic boundaries identified from surface features are important in the deep mesopelagic and bathypelagic. We present community analyses of discrete-depth samples of mesozooplankton and micronekton to full-ocean depth collected in the area where the Mid-Atlantic Ridge is crossed by the Subpolar Front. The results show that the distributional discontinuity associated with the front, which is strong near the surface, decreases with increasing depth. Both the frontal separation near the surface and the community convergence at increasing depths were clearer for mesozooplankton than for micronekton.

Biogeographic and palaeoclimatic relationships of the Middle Pliocene ichthyofauna of the Samoggia Torrent (Bologna, Italy)

New Middle Pliocene ichthyofauna (2.4-2.2 Ma) from central-eastern Italy (Samoggia Torrent, Bologna) are described. These ichthyolites were found in a rather thin laminated layer that was deposited after the 2.4 Ma climatic crisis. The origin of this deposit, in which 31 taxa have been classified, is to be related to anoxic events on a regional and, probably, supraregional scale. This ichthyofaunistic association, which consists of living genera, is characterized by a clearcut predominance of mesopelagic species. The palaeoclimatic characters of these ichthyofauna indicate subtropical-type waters, while from a palaeobiogeographic point of view there is a close relationship with the present-day Atlantic-Mediterranean bioprovince. The Samoggia deposit has yielded six taxa that are absent or only occasionally present in the Mediterranean: one of these, Spratelloides gracilis, is exclusive of the Indo-Pacific bioprovince.