P.M. Bagley

The fate of cetacean carcasses in the deep sea: observations on consumption rates and succession of scavenging species in the abyssal north-east Atlantic Ocean

The fate of cetacean carcasses in the deep sea was investigated using autonomous deep–sea lander vehicles incorporating time–lapse camera systems, fish and amphipod traps. Three lander deployments placed cetacean carcasses at depths of 4000 to 4800 m in the north–east Atlantic for periods of 36 h, 152 h and 276 h before being recovered. The photographic sequences revealed that carcasses were rapidly consumed by fish and invertebrate scavengers with removal rates ranging from 0.05 to 0.4 kg h-1. In the longest experiment the carcass was skeletonized within five days. In each deployment, approximately an hour after emplacement, the grenadier Coryphaenoides (Nematonurus) armatus and large numbers of lysianassid amphipods had arrived at the food–fall. The initially high numbers of grenadiers declined once the majority of the bait had been consumed and a variety of other fish and invertebrates were then observed, some taking up residence at the site. None of the fish species appeared to consume the carcass directly, but preyed upon amphipods instead. Funnel traps recovered with the carcass indicated a succession in the species composition of amphipods, with the specialist necrophages such as Paralicella spp. being replaced by more generalist feeders of the Orchomene species complex.

Photographic and acoustic tracking observations of the behaviour of the grenadier Coryphaenoides (Nematonurus) armatus, the eel Synaphobranchus bathybius, and other abyssal demersal fish in the North Atlantic Ocean

Using an autonomous free-fall vehicle (AU-DOS), observations were made of demersal fish attracted to baits and baited acoustic transmitters at two stations in the North Atlantic Ocean. A comparison was made between Station PAP (48°50′N; 16°30′W), 4800 m deep on the Porcupine Abyssal Plain which is relatively eutrophic, and Station MAP (31°N; 20°W), 4900 m deep on the Madeira Abyssal Plain, which is oligotrophic. Experiments were conducted during summer, in 1989 and 1990. Four species of fish were observed at Station MAP, the grenadier, Coryphaenoides (Nematonurus) armatus, the eel, Synaphobranchus bathybius, and the ophidiids Spectrunculus grandis, and Barathrites sp. At Station PAP, C. (N.) armatus and H. (S.) bathybius were attracted to bait on all deployments and only two other individuals of different species, probably ophidiids, were seen. The mean first grenadier arrival time was 30 and 138 min at Stations PAP and MAP, respectively. Mean first eel arrival time was 29 and 151 min at Stations PAP and MAP, respectively. Estimated population densities of fish were 167 grenadiers km-2 and 180 synaphobranchid eels km-2 at Station PAP and 8 grenadiers km-2 and 7 eels km-2 at Station MAP. Only the grenadier C. (N.) armatus definitely ingested transmitters, and this species dominated fish activity around the baits. Mean time of departure of grenadiers with transmitters in their stomachs across an acoustic horizon at 1000 m range was 371 and 488 min at Stations PAP and MAP, respectively. Grenadiers had a longer mean staying time at the food source at the more oligotrophic Station MAP (364 min) than at Station PAP (141 min). This corresponds with predictions of optimal foraging theory.

In situ comparison of activity in two deep-sea scavenging fishes occupying different depth zones

The activity of two scavenging deep–sea fishes occupying the same niche in overlapping depth zones were compared by in situ measurements of swimming speeds, tail–beat frequencies and by arrival time at baits. At 4800 m on the Porcupine Abyssal Plain, the grenadier Coryphaenoides (Nematonurus) armatus was the dominant scavenger, arriving at baits after 30 min, and swimming at relatively slow speeds of 0.17 body lengths (BL) sec-1. At 2500 m in the relatively food rich Porcupine Seabight both C. (N.) armatus and the blue–hake, Antimora rostrata, were attracted to bait, but A. rostrata was always the first to arrive and most of the bait was consumed before the C. (N.) armatus arrived. A. rostrata swam at mean speeds of 0.39 BL sec−1, similar to related shallow water species at equivalent temperatures. Observations on tail–beat frequency from video sequences confirmed the greater activity of A. rostrata. The data indicate that, given sufficient food supply, high pressure and low temperature do not limit activity levels of demersal deep–sea fishes. Low activity of C. (N.) armatus is an adaptation to poor food supply in the abyss, where these fishes dominate, but prevents it competing with the more active A. rostrata in shallower depths.

Liparid and macrourid fishes of the hadal zone: in situ observations of activity and feeding behaviour

Using baited camera landers, the first images of living fishes were recorded in the hadal zone (6000–11 000 m) in the Pacific Ocean. The widespread abyssal macrourid Coryphaenoides yaquinae was observed at a new depth record of approximately 7000 m in the Japan Trench. Two endemic species of liparid were observed at similar depths: Pseudoliparis amblystomopsis in the Japan Trench and Notoliparis kermadecensis in the Kermadec Trench. From these observations, we have documented swimming and feeding behaviour of these species and derived the first estimates of hadal fish abundance. The liparids intercepted bait within 100–200 min but were observed to preferentially feed on scavenging amphipods. Notoliparis kermadecensis act as top predators in the hadal food web, exhibiting up to nine suction-feeding events per minute. Both species showed distinctive swimming gaits: P. amblystomopsis (mean length 22.5 cm) displayed a mean tail-beat frequency of 0.47 Hz and mean caudal : pectoral frequency ratio of 0.76, whereas N. kermadecensis (mean length 31.5 cm) displayed respective values of 1.04 and 2.08 Hz. Despite living at extreme depths, these endemic liparids exhibit similar activity levels compared with shallow-water liparids.

Estimating the abundance of Patagonian toothfish Dissostichus eleginoides using baited cameras: a preliminary study

The Patagonian toothfish Dissostichus eleginoides has been the object of a rapidly expanding longline fishery in the Southern Ocean. Little is known about the biology of D. eleginoides and traditional methods of estimating stock size using trawling techniques have proved ineffective because the adult fish are found in deep waters on the continental slope at depths of 700–2500 m. During September 1997, a preliminary study was undertaken using arrival times at an autonomous baited camera vehicle, the Aberdeen University Deep Ocean Submersible (AUDOS), to estimate the abundance and size of toothfish in waters around South Georgia (SG) and the Falkland Islands (FI). These are the first attempts at estimating the abundance of toothfish that are independent of catch data from the commercial fishery.

A Large Aggregation of Liparids at 7703 meters and a Reappraisal of the Abundance and Diversity of Hadal Fish

Few biological studies have investigated the hadal depths within oceanic trenches that plummet from 6000 meters (m) to the full ocean depth of almost 11,000 m. Here we present the deepest known in situ observations of fish: a hadal snailfish, Pseudoliparis amblystomopsis (Andriashev 1955), from 7703-m deep in the Japan Trench, which was obtained using a baited video lander. The maximum number of fish we observed was unexpectedly higher than trawl catch records of any known hadal fish. We describe changes in fish abundance and associated behaviors over time, including feeding, resting, and swimming. In light of these new observations, we reappraise the occurrence and diversity records of hadal fishes that have been constructed from fragmentary and often misleading information derived from historical explorations and global data sets. This reappraisal suggests that hadal fish diversity may be lower—although some hadal fish species may attain much larger populations—than previously thought.

High swimming and metabolic activity in the deep-sea eel Synaphobranchus kaupii revealed by integrated in situ and in vitro measurements.

Several complementary studies were undertaken on a single species of deep‐sea fish (the eel Synaphobranchus kaupii) within a small temporal and spatial range. In situ experiments on swimming and foraging behaviour, muscle performance, and metabolic rate were performed in the Porcupine Seabight, northeast Atlantic, alongside measurements of temperature and current regime. Deep‐water trawling was used to collect eels for studies of animal distribution and for anatomical and biochemical analyses, including white muscle citrate synthase (CS), lactate dehydrogenase (LDH), malate dehydrogenase (MDH), and pyruvate kinase (PK) activities. Synaphobranchus kaupii demonstrated whole‐animal swimming speeds similar to those of other active deep‐sea fish such as Antimora rostrata. Metabolic rates were an order of magnitude higher (31.6 mL kg−1 h−1) than those recorded in other deep‐sea scavenging fish. Activities of CS, LDH, MDH, and PK were higher than expected, and all scaled negatively with body mass, indicating a general decrease in muscle energy supply with fish growth. Despite this apparent constraint, observed in situ burst or routine swimming performances scaled in a similar fashion to other studied species. The higher‐than‐expected metabolic rates and activity levels, and the unusual scaling relationships of both aerobic and anaerobic metabolism enzymes in white muscle, probably reflect the changes in habitat and feeding ecology experienced during ontogeny in this bathyal species.

Measurement of in situ oxygen consumption of deep-sea fish using an autonomous lander vehicle

Conventional laboratory studies of deep-sea fish metabolism are not possible as these fish are typically killed during recovery to the surface. As these species are important members of deep-sea communities, the lack of these data represents a significant limitation to our understanding of the functioning of this ecosystem. An autonomous fish respirometer vehicle was developed in order to measure the oxygen consumption of deep-sea fish in situ. This new lander allows measurements to be made without handling or stressing the animals and without the logistical problems and great cost of submersible operations. The design, operation, and measurement methodology are described and preliminary data for Coryphaenoides armatus at 4000 m are presented. These Atlantic data appear to confirm the low metabolic rate measured in this species when compared to other gadid species.

In situ investigation of burst swimming and muscle performance in the deep-sea fish Antimora rostrata (Gunther, 1878)

The few existing measurements of deep-sea fish physiology consistently indicate reduced basal metabolism and metabolic power. A possible explanation for this is the reduction in selective pressure for burst activity capacity due to a reduction in the frequency and duration of predator-prey interactions in the sparsely distributed fish community and continuous darkness. Video recordings of stimulated fast-starts in deep-sea fish were obtained by a lander vehicle and analysed to give the swimming velocities, accelerations, and inertial power requirements of fast-start swimming in Antimora rostrata. With a mean peak velocity of 0.7 m s(-1), and white muscle power output of only 17.0 W kg(-1). A. rostrata is a slow moving fish, but no slower than shallow-water fishes at the same temperature.

Scavenging by megabenthos and demersal fish on the South Georgia slope

The scavenging megafauna of the South Georgia and Shag Rocks slope in the south-west Atlantic (625–1519 m) were investigated using autonomous baited camera systems. Two surveys were conducted: the first in 1997 (13 deployments) used a conventional 35 mm stills camera with a 200 J flash, whilst the second in 2000 (15 deployments) used low-light digital video cameras. The scavenging community responded rapidly to the arrival of bait on the sea floor and was dominated by stone crabs (Lithodidae) and toothfish (Dissostichus eleginoides). Stone crabs took up residence around the bait until it was consumed, with a maximum number of 108 in the field of view after four hours. The most frequently observed crab species was Paralomis formosa. Paralomis spinosissima, Neolithodes diomedea and Lithodes sp., were also observed. Toothfish were the most frequently observed scavenging fish and were seen during all but one deployment, typically making brief visits (1–2 min) to the bait, but appeared startled by the flash in the 1997 survey. Labriform swimming (sculling with the pectoral fins) was the principal form of locomotion in toothfish (0.22 body lengths (BL) sec−1), but they were capable of more rapid sub-carangiform (using caudal trunk and fin) motion (3 BL sec−1) when startled. Other scavenging fish observed included the blue-hake Antimora rostrata, grenadiers (Macrourus spp.), skates, liparids and zoarcids.