The diving behaviour of brooding king penguins (<Emphasis Type="Italic">Aptenodytes patagonicus</Emphasis>) from the Falkland Islands: variation in dive profiles and synchronous underwater swimming provide new insights into their foraging strategies

The diving behaviour of king penguins (Aptenodytes patagonicus) was studied on the Falkland Islands, where a small population (ca. 300 fledglings year^–1) is located at the geographical limit of their breeding range. King penguins rearing newly hatched chicks were equipped with time-depth recorders before leaving for sea. In total, 20,175 dives >3 m were recorded from 12 birds during 15 foraging trips with a mean duration of 5.7±2.3 days. The majority of the trips was directed up to 500 km to the northeast of the breeding colony in slope waters of, and oceanic waters beyond, the Patagonian shelf. Mean time spent underwater accounted for 42±9% of the foraging trip. Mean dive depth achieved was 55±16 m; maximum dive depth recorded was 343 m. Mean dive duration was 159±25 s; maximum dive duration was 480 s. The mean vertical distance covered was 140±65 km trip^–1; and on average birds covered 25 km day^–1. Synchronous diving behaviour was observed in two birds for a period of about 24 h after leaving the colony. Dive depth correlated positively with: (1) light intensity, (2) dive duration and (3) vertical velocities, thus confirming previous findings obtained from conspecifics at other breeding sites and indicating comparable diving behaviour. However, separation of dives according to their profile—V-, U-, or W-shaped—revealed significant differences between certain dive parameters. For a given depth range, bottom time was longer and vertical velocities higher in W-dives than in U-dives. This, together with a higher number of W-dives at dawn and dusk, suggests that foraging is more effective during W-dives than U-dives, and during twilight. These findings imply that king penguins have to make more complex decisions, individually and socially, on the performance of the subsequent dive than previously thought.Communicated by O. Kinne, Oldendorf/Luhe     

Ein fotoschlitten für biologische und geologische kartierungen des meeresbodens

A photosledge is described; it is 4 m long and 2 m wide and can be used also on an uneven sea bottom. In its foremost part a camera is mounted in a pressure sphere which has been tested for water depths down to 6,000 m. The camera system can hold 90 m of film; with an exposure size of 24x24 mm this accounts for 3,600 exposures. The time between 2 exposures may be varied from 1 to 10 sec and this allows to cover a distance of 2 to 20 nautical miles at a speed of 2 knots within 1 single mission. The photographic angle may be changed from vertical to horizontal. Best results have been obtained with angles of 30 and 45 degrees from the vertical. The electronic flash, with a power of 60 Wsec, is mounted below the camera. Successful work has been conducted in the North Sea and the Baltic Sea at depths between 40 and 50 m, on the continental slope off Portugal between depths of 140 and 240m, and on top of Great Meteor Seamount at depths between 280 and 350 m.     

Orientation ofRhizophagus grandis (Coleoptera: Rhizophagidae) to oxygenated monoterpenes in a species-specific predator-prey relationship

Summary The predator,Rhizophagus grandis, is linked to its specific prey,Dendroctonus micans, by semiochemical signals regulating oviposition and long range orientation. A mixture of simple oxygenated monoterpenes [(–)—fenchone, (–)—pinocamphone, rac. camphor, terpinene-4-ol, borneol, fenchol and verbenone], identified from the frass ofD. micans has been found to be extremely active in a flight windtunnel as a long—range attractant for the predator. The mixture elicited 84 % of the response to larval frass of the prey. Excluding pinocamphone from the mixture did not influence its activity. Also, changing the absolute configurations of some of the components (fenchol, terpinene-4-ol and borneol) did not influence the predators' response. However, the addition of (–)—-terpineol increased the attractivity of the synthetic blend to almost that of larval frass of the prey. Ecological implications of the identified semiochemicals and their use in pest management are discussed.     

A new towed underwater camera system for wide-range benthic surveys

Following a short review of previous underwater camera systems, a new combined underwater camera system, consisting of a remote controlled TV- and photocamera mounted on a ship-towed steel-tubing sled, is described. The method of using this apparatus for wide-range surveys of the marine littoral benthos, and the techniques used to evaluate and quantify the observations are presented. On the basis of first results obtained in the North Adriatic Sea, the efficiency of the new system is compared with various other instruments and techniques. Advantages and disadvantages, possible improvements, and future prospects are discussed.     

Field observations on the emergence rhythm of the Norway lobsterNephrops norvegicus,using different methods

The emergence ofNephrops norvegicus (L.) from their burrows was studied on several fishing grounds around Scotland at 28 to 62 m depth. Various methods were used: sampling by trawl and baited creels, observations by television and time-lapse cameras. Peak emergence generally occurred around dusk and dawn. In shallow water (30 m) there was also a significant level of activity during the intervening night period. Day/night variations in the size composition of catches suggest that relatively more time is spent out of the burrow with increasing size. There was good agreement between the various methods employed, suggesting that trawl and camera avoidance has little influence on the pattern of emergence observed. The use of bait in some experiments confirmed that emergence is essentially a feeding activity.     

A Reliability and Risk Analysis System for Multipurpose Reservoir Operation

This paper proposes a reliability analysis system, which can be widely applied to the cases in which a reservoir is operated to meet several purposes such as flood control, energy generation, irrigation, domestic and industrial water supply, etc. The presented system has a structure of three levels.1. Decision-making.2. Tradeoff analysis.3. The third level that mainly consists of three subsystems:i – The reservoir flood risk analysis subsystem by flood control simulation.ii – The benefit promotion subsystem of reservoir operation using Stochastic Dynamic Programming. In this subsystem, the Lagrange multipliers are introduced into the objective function to take into account the water supply failures. This method guarantees that each run of the SDP will necessarily derive a non-inferior policy for reservoir operation.iii – The reservoir operation simulation subsystem to derive the performance indices associated with the reservoir operation policies. With the input and feedback between the second level and the subsystems of the third level, a great deal of efficient operation policies and the associated performance indices can be obtained. Then the tradeoff relationships between different performance indices can be derived for the decision makers. With application to Yudong Reservoir in Yunnan province of China, the presented analysis system is practically tested.     

Assessment of herbaceous plant habitats in water-constrained environments: predicting indirect effects with fuzzy logic

Herbaceous plant production plays a key role in determining the function of rangeland ecosystems in the semi-arid and Mediterranean regions. Therefore, assessment of herbaceous plant habitats is important for understanding the ecosystem functioning in these regions and for applied purposes, such as range management and land evaluation. This paper presents a model to assess herbaceous plant habitats in a basaltic stony environment in a Mediterranean region. The model is based on geographic information systems (GIS), remote sensing and fuzzy logic, while four indirect variables, which represent major characteristics of herbaceous habitats, are modeled: rock cover fraction; wetness index (WI); soil depth; and slope orientation (aspect). A linear unmixing model was used to measure rock cover on a per pixel basis using a Landsat TM summer image. The wetness index and local aspect were determined from digital elevation data with 25 m × 25 m pixel resolution, while soil data were gathered in a field survey. The modeling approach adopted here is process-based and assumes that water availability plays a crucial role in determining herbaceous plant production in Mediterranean and semi-arid environments. The model rules are based on fuzzy logic and are written based on the hypothesized water requirements of the herbaceous vegetation. The results show that on a polygon basis there is positive agreement between the model proposed here and previous mapping of the herbaceous habitats carried out in the field using traditional methods. Intrapolygon tests show that the use of a continuous raster data model and fuzzy logic principles provide an added value to traditional mapping. Moreover, herbaceous biomass measurements at two time intervals—mid- and peak winter season—corresponded with the habitat assessment predictions achieved using a new scenario that is proposed in this research. This scenario suggests that rockiness increases herbaceous production on south-facing slopes, while in other slope aspects the rock cover has lower impact on herbaceous growth. Due to its simplicity, the model suggested here can be used by planners and managers, to adjust range activities over large areas. The process-based approach should allow adaptation of the model to other regions more effectively than models that were formulated on a purely empirical basis. The model could also be used to study the relationship between water availability and ecosystem productivity on a regional scale.     

Functional effects of the hadal sea cucumber <Emphasis Type="Italic">Elpidia atakama</Emphasis> (Echinodermata: Holothuroidea,Elasipodida) reflect small-scale patterns of resource availability

Holothuroidea represent the dominant benthic megafauna in hadal trenches (~6,000–11,000 m), but little is known about their behaviour and functional role at such depths. Using a time-lapse camera at 8,074 m in the Peru–Chile Trench (SE Pacific Ocean), we provide the first in situ observations of locomotory activity for the elasipodid holothurian Elpidia atakama Belyaev in Shirshov Inst Oceanol 92:326–367, (1971). Time-lapse sequences reveal ‘run and mill’ behaviour whereby bouts of feeding activity are interspersed by periods of locomotion. Over the total observation period (20 h 25 min), we observed a mean (±SD) locomotion speed of 7.0 ± 5.7 BL h⁻¹, but this increased to 10.9 ± 7.2 BL h⁻¹ during active relocation and reduced to 4.8 ± 2.9 BL h⁻¹ during feeding. These observations show E. atakama translocates and processes sediment at rates comparable to shallower species despite extreme hydrostatic pressure and remoteness from surface-derived food.     

Complexity of coral interactions: Influence of time,location of interaction and epifauna

The potential role of competition for space in a community depends on the arrangement of interaction relationships. A survey (255 m²) of the interactions between corals (Scleractinia) on a Caribbean reef (depth 10–30 m) indicated the outcome of 17–35% of the aggressive and defensive interactions to be unpredictable. Experiments on the reef (depth 7–13 m) with pairs of interacting corals — Madracis mirabilis (Duchassing & Michelotti), Agaricia agaricites (L), Montastrea annularis (Ellis & Solander), Eusmilia fastigiata (Pallas) — showed that, after the initial contest through extracoelenteric digestion, there are at least two additional processes which can result in a reversal of dominance: interference by epifauna and sweeper tentacle development. Moreover, the impact of extracoelenteric digestion and the extent of sweeper tentacle development varied over the surface of the corals. Employing laboratory and field experiments to distinguish between the impact of extracoelenteric digestion, epifauna behaviour and sweeper tentacles, we show the three processes combined to explain the coral interaction process in toto. The outcome of the interaction process on the reef depends on numerous, partly unpredictable, variables, including mode of contact and effects of position. Consequently, patterns of community organization resulting from spatial competition will be slow to emerge and easily erased prematurely by disturbances.     

Prodissoconch morphology is environmentally modified in the brooding bivalve Lasaea subviridis

Embryonic shell sculpture of intertidal and laboratory brooded individuals of the direct developer Lasaea subviridis was studied using scanning electron microscopy. Intertidally brooded individuals develop a distinct prodissoconch I (PI) and prodissoconch II (PII) of unusual morphologies. The PI is relatively small (50–100 m in length), oval in outline, lacks a radial sculpture and is restricted to the initial pitted zone of the prejuvenile shell. A larger PII is formed (510–680 m in length) and it is dominated by pronouced commarginal striae but also includes a belt of radial ridges extending from the PI/PII boundary. Laboratory brooded individuals kept constantly submerged do not develop the pronounced commarginal striae characteristic of PII. This implies that PII formation in L. subviridis is not a direct result of the brooding habit, and can be modified by environmental factors, which possibly include low-tide exposure. There is no single feature of external prodissoconch morphology that unambiguously indicates a direct development mode combined with mantle cavity brood protection. General guidelines are presented to help recognize this developmental mode, based on prodissoconch morphology. In comparing prodissoconch morphologies of brooding bivalves, the habitats as well as the phylogenies of species should be considered, especially in comparisons of intertidal and subtidal species.     

The poison gland of the antMyrmicaria eumenoides and its role in recruitment communication

Summary The antMyrmicaria eumenoides is a significant arthropod predator. For rapid attraction of large numbers of nestmates to newly discovered food sources the ants use an efficient recruitment communication system based on the poison gland secretion. Workers exhibit age-based division of labour. Young workers perform brood-care; their poison gland reservoir develops and reaches its final size of 0.5 µl at an age of 6 weeks, when they become foragers. The secretion deposited during combat with enemies or prey is composed of equal amounts of both a high volatile and a low volatile fraction. Within the high volatile fraction (+)—limonene is the main component (97%) and is the only olfactory trigger to alert ants in the vicinity and to recruit them to places of combat, where they assemble. Ants respond to synthetic (+)—limonene in exactly the same way as to the poison gland secretion when applied at the same airborne concentrations. Further components of the high volatile fraction are four additional monoterpene hydrocarbons and hexanoic nitrile. The high volatile and the low volatile fraction of the poison gland secretion each have dual functions: The low volatiles, of which the main component is an alkaloid, serve as a fixative and extend the effective period of the limonene signal by modifying its evaporation kinetics. On the other hand the high volatile recruitment signal (+)—limonene is also the solvent for the alkaloid and enhances its spreading on the surface of the cuticle of arthropod enemies or prey.     

Vertical distribution of macroplankton at the Charlie-Gibbs Fracture Zone (North Atlantic), as observed from the manned submersible “Mir-1”

Direct observations of the plankton vertical distribution performed from a manned submersible were made in the Charlie-Gibbs Fracture Zone (Atlantic Ocean). Plankton was counted using a standard 3-m² frame in the depth layer from 170 to 4361 m and plotted for every 50 m. The most abundant components of the macroplankton were pelagic shrimps, chaetognaths and gelatinous animals, with peak of densities corresponding to the main pycnocline. Mucous houses of appendicularians were abundant at 150 m above the seabed—up to 0.07–0.09 m^–3, compared to 0–0.006 m^–3 in the upper layers. The new observations confirm that the near-bottom peak of appendicularian abundance is characteristic of at least the entire Central Atlantic, and it is not associated with certain biotopes on the bottom.Communicated by O. Kinne, Oldendorf/Luhe

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Induction of trypsinogen secretion in herring larvae (Clupea harengus)

Mechanisms initiating trypsinogen secretion were studied in laboratory reared herring larvae (Clupea harengus L.) exposed to physical and chemical stimuli. Pancreatic secretion of trypsinogen was quantified for each stimulus type as the increase above pre-stimulus level of intestinal trypsin content. Larval prey types were: nauplii, copepodites or adult Acartia tonsa, small polystyrene spheres (diameter 94 m), small (diameter 79 m) or large (diameter 170 m) polystyrene-latex spheres. Intestinal trypsin content can be expressed as a function of two variables: meal size and content of pancreatic trypsinogen. Trypsinogen secretion increases with different prey items in the order: small spheres, nauplii and copepodites. Larvae which eat large spheres secrete more enzyme than if fed small spheres but trypsinogen secretion is similar in fish larvae fed copepodites and large spheres. The fact that the size of non-biodegradable particles exerts a major control over trypsinogen secretion suggests neural — as opposed to chemically mediated — initiation of secretion. A cephalic phase of secretory stimulation could not be demonstrated during swallowing of copepods or exposure for 2 to 3 h to compounds which leak from live copepodites. As cephalic and gastric phases of secretory stimulation are absent, initiation of trypsinogen secretion must take place in the intestine. Larval herring retain trypsin in the intestine. Ca. 4.5 h after a meal, 3/4 of the enzyme is located in the intestinal fluid, presumably available for hydrolysis of subsequent meals, and the high proportion (ca. 25%) of the pancreatic trypsinogen content which is secreted for copepodite prey may thus not be energetically wasteful for the larvae.     

Life in a warm deep sea: routine activity and burst swimming performance of the shrimp<Emphasis Type="Italic"> Acanthephyra eximia</Emphasis> in the abyssal Mediterranean

Measurements of routine swimming speed, tail-flip escape responses, and oxygen consumptions were made of the deep-sea shrimp Acanthephyra eximia using autonomous landers in the Rhodos Basin at depths of up to 4,400 m and temperatures of 13–14.5°C. Routine swimming speeds at 4,200 m averaged 0.18 m s^–1 or 3.09 body lengths s^–1, approximately double those of functionally similar oceanic scavengers. During escape responses peak accelerations of 23 m s^–2 or 630.6 body lengths s^–2 were recorded, with animals reaching speeds of 1.61 m s^–1 or 34.8 body lengths s^–2. When compared to shallow-water decapods at similar temperatures these values are low for a lightly calcified shrimp such as A. eximia despite a maximum muscle mass specific power output of 90.0 W kg^–1. A preliminary oxygen consumption measurement indicated similar rates to those of oceanic crustacean scavengers and shallower-living Mediterranean crustaceans once size and temperature had been taken into account. These animals appear to have high routine swimming speeds but low burst muscle performances. This suite of traits can be accounted for by high competition for limited resources in the eastern Mediterranean, but low selective pressure for burst swimming due to reductions in predator pressure.Communicated by J.P. Thorpe, Port Erin     

A model for the larval migration of the Japanese eel: roles of the trade winds and salinity front

A model that helps explain the mysterious long-distance migration of the Japanese eel (Anguilla japonica) is presented, based on oceanographic observations, satellite buoy drift experiments, and samplings of eel larvae taken in 1991. The trajectory of a 150 m depth buoy relased in the spawning area strongly suggests that A. japonica larvae spawned just south of the salinity front are transported westward by the North Equatorial Current (NEC). The larvae are then thought to be entrained into the Mindanao Current flowing southward along the Philippine Islands where A. japonica juveniles are scarcely distributed. These controversial results lead to the assumption that eel larvae are transferred from the NEC to the northward flowing Kuroshio, which distributes the eel larvae to the growth habitats of eastern Asia. In this eel larvae transfer model, a northward Ekman transport caused by trade winds plays an important role in explaining the wind-induced northward shift of the larvae together with the onset of diel vertical migration. Assuming that leptocephali greater than 20 mm initiate the vertical migration, a westward wind velocity greater than 5 to 10 m s^-1 should be high enough to diminish the southward current velocity. When the physical and geophysical conditions — such as the salinity front for spawning activity, the water tunnel for westward larval transport, the Ekman transport by the trade wind for transfer of the larvae from the NEC to the Kuroshio, and the strong velocity of the Kuroshio for rapid transport to growth habitats — are well matched with the timing of the onset of vertical migration, large-scale eel migration could result.     

Locomotory activity and behaviour of the Antarctic teleostNotothenia coriiceps

The activity and behaviour of a free-living Antarctic fish,Notothenia coriiceps Richardson (formerlyN. neglecta), was investigated using a high-sensitivity, underwater TV camera at Signy Island, South Orkney Islands. Detailed observations of the 33 cm TL (total length) fish were made over a period of 6 d in austral summer (February 1992), for a total 69.5 h. Natural light at 2.5 m depth allowed viewing from 1 h before sunrise to 1 h after sunset. The fish stayed in a territory within 3 m of a small cave for >98% of the time, and made between 1 to 148 swims d^-1, of which 92.5% were brief (<15 s) feeding attempts. On average, 1.7% of each day was engaged in locomotion, including 1.2% swimming and 0.5% manoeuvring. Swimming was generally slow, at <2 body lengths s^-1, and labriform and subcarangiform modes were used alternately or in combination. Activity level (swims or displays per hour) was unaffected by tide, but was lower for 3 d when a wind speed >16 knots prevailed indicating that large waves reduced activity. A suspected diurnal activity rhythm was not statistically significant. The fish is an ambush-predator, and it took most of its prey from the water column but some off macroalgae or the seabed. Ventilation rate was slightly higher after activity, and peaked after an encounter with anotherN. coriiceps.     

An EM algorithm for capture-recapture estimation

Analysis of capture—recapture data often involves maximizing a complex likelihood function with many unknown parameters. Statistical inference based on selection of a proper model depends on successful attainment of this maximum. An EM algorithm is developed for obtaining maximum likelihood estimates of capture and survival probabilities conditional on first capture from standard capture—recapture data. The algorithm does not require the use of numerical derivatives which may improve precision and stability relative to other estimation schemes. The asymptotic covariance matrix of the estimated parameters can be obtained using the supplemented EM algorithm. The EM algorithm is compared to a more traditional Newton-Raphson algorithm with both a simulated and a real dataset. The two algorithms result in the same parameter estimates, but Newton-Raphson variance estimates depend on a numerically estimated Hessian matrix that is sensitive to step size choice.     

Light harvesting by wavelength transformation in a symbiotic coral of the Red Sea twilight zone

We report an extraordinary depth range for Leptoseris fragilis (Milne Edwards and Haime), a reef building coral of the Red Sea living in cytosymbiosis with zooxanthellae. The coral harbours an as yet unknown pigment system. We suggest that the heterotrophic host — the coral — provides its photoautotrophic symbionts with additional light. The supplementary light is provided by host pigments which transform light of short wavelengths into suitable wavelengths for photosynthesis, thus amplifying and increasing the transfer of photoassimilates from the zooxanthellae to the host.     

Spoil disposal

Summary The colliery spoil problem in existing mining areas requires the same attention as that in the new coalfields. The Yorkshire coalfield has particular problems since so much land near collieries has already been tipped on. Suggested short-term and long-term solutions are outlined. Some will be successful, others may not.The Commission on Energy and the Environment has made many recommendations to improve the existing situation in the coalfields. How many of these will the Government support? The National Coal Board promised a high standard of environmental treatment at the Belvoir public inquiry. Will these high standards be reflected in developments on the existing coalfields?Effective day-to-day management can have beneficial results out of all proportion to expenditure. All collieries should match up to the high standards achieved at the better ones. Is there any reason why lagoons should be features of colliery spoil disposal for the next decade? Can colliery spoil be used more extensively as an alternative material in construction? The National Coal Board and the local authorities need a strategic plan for coal development including spoil disposal and in detail need an agreed plan for short-term spoil disposal at each colliery.Commitment and research is needed to look into methods of backstowage — which tackles the problem at source. Can other European experience help in this matter? Fluidised-bed technology may lead to a reduction in spoil. Maximum use will have to be made of local tipping opportunities, whether quarry, voids, opencast coal sites, new land forms or upgrading agricultural land — but they won't happen on their own. Effort and determination will be needed. Long-distance movement of spoil appears expensive. It cannot be financed under the NCB's present system of accounting and certainly not financed by local authorities. Costs may need to be met nationally. The financing of major spoil solutions may well be the major problem in making and progress in any direction — but a national resource such as coal should be developed through a national approach to overall costs.     

A criterion validity test of the contingent valuation method: comparing hypothetical and actual voting behavior for a public referendum

This study pursues external validation of contingent valuation by comparing survey results with the voting outcome of a Corvallis, Oregon, referendum to fund a riverfront improvement project through increased property taxes. Survey respondents hypothetically make a voting decision—with no financial consequences—on the upcoming referendum. The survey sample consists of respondents verified to have voted in the election. We use available precinct-level election data to compare the proportion of “yes” survey and referendum votes as well as estimate voting models and mean willingness to pay (WTP) based on the two sets of data. We find that survey responses match the actual voting outcome and WTP estimates based on the two are not statistically different. Contrary to similar studies, our statistical results do not depend on re-coding the majority of “undecided” survey responses to “no.” Furthermore, such a re-coding of responses may be inappropriate for our data set.