Stereophotography for the field biologist: measurement of lengths and three-dimensional positions of free-swimming sharks

A stereophotographic technique for determining size and relative position of free-swimming sharks is described and illustrated for schooling scalloped hammerhead sharks (Sphyrna lewini). The method yields total length and nearest-neighbor, interindividual distance; and each of these dimensions is expressed as a function of the shark's distance into a school on the vertical and horizontal planes. Stereopairs of photographs were taken by an aligned, beam-mounted pair of cameras (Nikonos III). The scale to determine the length of a shark from the paired photographic images was obtained from the horizontal displacement between the images. Displacement was correlated with optical axis separation from photographs of a scaled staff at known distances from the camera. Image dimensions on the photographs were measured by projecting a scale onto the stage through a camera lucida. The precision of repeated measurements of a 50 cm section of a scaled staff at increasing distances from the cameras of 2, 4, and 8 m was ±5.0%. Lengths of the sharks ranged from 109 to 371 cm, with a median of 178 cm, for 3 offshore sites in the Gulf of California during July and August 1979. At one site, El Bajo Gorda, lengths increased with both distance from the camera and distance into the group; in contrast, the interindividual distances (head-to-head) did not vary with distance into the group and possessed a median of 232 cm.     

The three-dimensional structure of airborne bird flocks

Summary The three-dimensional structure of flocks of dunlin, Calidris alpina, and starlings, Sturnus vulgaris, was studied while birds were in transit between feeding, loafing and roosting sites. A technique was developed that uses standard photogrammetric methods to determine the three-coordinate position of birds in flocks from stereoscopic pairs of simultaneously exposed photographs. A comparison of nearest neighbour distances indicates that dunlin have a tighter, more compact flock structure than do starlings (Fig. 2; Table 2). Analysis of interbird angles in both the vertical and horizontal planes indicates that each dunlin's nearest neighbour is most likely to be behind and below it. This spatial structure results in areas in which few nearest neighbours occur (e.g., immediately in front and below) (Fig. 3). Flight speeds during transit flights are also presented (Table 4). The spatial structure and behaviour of dunlin and starling flocks appear to be very similar to the structure and behaviour of schools of fish.     

Foraging behavior in early Atlantic cod larvae (Gadus morhua) feeding on a protozoan (Balanionsp.) and a copepod nauplius (Pseudodiaptomussp.)

Food limitation is likely to be a source of mortality for fish larvae in the first few weeks after hatching. In the laboratory, we analyzed all aspects of foraging in cod larvae (Gadus morhua Linnaeus) from 5 to 20 d post-hatching using protozoa (Balanion sp.) and copepod nauplii (Pseudodiaptomus sp.) as prey. A camera acquisition system with two orthogonal cameras and a digital image analysis program was used to observe patterns of foraging. Digitization provided three-dimensional speeds, distances, and angles for each foraging event, and determined prey and fish larval head and tail positions. Larval cod swimming speeds, perception distances, angles, and volumes increased with larval fish size. Larval cod swam in a series of short intense bursts interspersed with slower gliding sequences. In 94% of all foraging events prey items were perceived during glides. Larval cod foraging has three possible outcomes: unsuccessful attacks, aborted attacks, and successful attacks. The percentage of successful attacks increased with fish size. In all larval fish size classes, successful attacks had smaller attack distances and faster attack speeds than unsuccessful attacks. Among prey items slowly swimming protozoans were the preferred food of first-feeding cod larvae; larger larvae had higher swimming speeds and captured larger, faster copepod nauplii. Protozoans may be an important prey item for first-feeding larvae providing essential resources for growth to a size at which copepod nauplii are captured. Received: 20 April 1999 / Accepted: 12 January 2000     

Daytime surface swarms of the euphausiid Thysanoessa inermis off the west coast of Hokkaido,northern Japan

Biological characteristics of the daytime swarming of the euphausiid Thysanoessa inermis (Krøyer) were examined off the west coast of Hokkaido in the northeast Sea of Japan between 1982 and 1987. Swarms were composed exclusively of fully mature individuals; males possessed spermatophores at the ejaculatory ducts and females had attached spermatophores at the thelycum. Females predominated in most swarms, while males or nearly balanced swarms were infrequently encountered. Fully mature females occurred in the late swarming season, usually mid-April. Examination of stomachs indicated that feeding activity was low during the daytime surface swarming behavior. The body sizes of swarming T. inermis occurring around Yagishiri Island were significantly larger than those found simultaneously off the Shakotan Peninsula. This suggests that the euphausiid populations of the two areas are different.     

Effect of light intensity on activity and food-searching of larval herring,Clupea harengus: a laboratory study

Larvae of Clupea harengus were reared from spawning herring caught in March 1982 and 1983 in the Firth of Clyde, Scotland. An infra0red observation technique was used to record the behaviour of larval herring both in shallow dishes using a top view and in a tank 2 m deep using a side view. The amount of time larvae spent swimming, which was minimum in complete darkness, increased with increasing light intensity and as the larvae grew. Maximum swimming speeds of feeding larvae were recorded at light intensities between 10 and 100 lux. The presence of food organisms (Artemia sp., Brazilian strain) at light intensities below the feeding threshold (0.1 lux) caused an increase in the proportion of time spent active, but light intensities above the threshold had different effects, depending on developmental stage: larvae of 12 mm increased swimming speed, but 21 mm larvae decreased speed. In the 2 m deep tank in darkness, larvae displayed inactive periods wherein they sank head first, interspersed with periods of upward swimming. As light intensity increased, vertical swimming was replaced by horizontal swimming. These results are discussed with reference to food searching and vertical migration of larval herring in the sea.     

Daytime surface swarming of Euphausia pacifica (Crustacea: Euphausiacea) in the Sanriku coastal waters of northeastern Japan

Biological characteristics of daytime surface swarming of Euphausia pacifica were investigated. Swarms in the Sanriku coastal waters consisted almost exclusively of mature individuals and seemed to be related to breeding behavior. Although maturation of ovarian eggs occurred both at the beginning and the end of the swarming period, females with attached spermatophore were most frequent at the end of the period. Since stomachs of swarming individuals were nearly empty throughout this period, swarming behavior was probably not related to feeding. Swarming individuals did not seem to be transported to the surface by vertical water movement caused by tidal streams, because the daily catch per boat did not differ between spring and neap tides.     

Ultrastructural study of the exoskeleton of the estuarine crab Rhithropanopeus harrisii: Effect of the insect growth regulator Dimilin® (diflubenzuron) on the formation of the larval cuticle

Ultrastructure of larval cuticle during the molt cycle of the estuarine crab Rhitropanopeus harrisii (Gould) (Crustacea: Brachyura) was studied in control larvae as well as in larvae exposed to 10 ppb of the insect growth regulator Dimilin^® (diflubenzuron). First zoeal larvae were used as test organisms. It has earlier been shown that 10 ppb Dimilin is lethal to zoeal larvae of R. harrisii, and nearly all exposed larvae died during molting to the next stage (Christiansen et al., 1978). Distinct differences in structure of the cuticle were found between the two groups of larvae. Both endocuticle and exocuticle appear to be deformed in Dimilin-treated larvae, whereas formation of epicuticle did not seem to be affected. The results indicate that Dimilin probably inhibits chitin synthesis in crab larvae as shown earlier by several authors for insect larvae.     

Effects of prey escape ability,flow speed,and predator feeding mode on zooplankton capture by barnacles

Experimental studies of feeding on zooplankton often involve the use of non-evasive Artemia spp. to represent zooplanktonic prey. Some zooplankton, however, such as copepods, are potentially evasive due to possession of effective predator-avoidance mechanisms such as high-speed escape swimming. In the present study, we compared the efficiencies with which non-evasive (A. salina) and evasive (copepods) zooplankton were captured by a sessile, suspension feeder, the coral-inhabiting barnacle Nobia grandis (Crustacea, Cirripedia). N. grandis specimens and zooplankton used in the present study were collected near Eilat, Israel in 1993. The effect of different flow speeds (from 0 to 14 cm s^-1) on captures of the two preys was also investigated. Additionally, we examined the effect of a flow-induced barnacle behavioral switch from active to passive suspension feeding, on zooplankton capture. Two video cameras were used to make close-up, three dimensional recordings of predator-prey encounters in a computer-controlled flow tank. Frame-by-frame video analysis revealed a highly significant difference (P< 0.001) in the efficiency with which A. salina and copepods were caught (A. salina being much more readily captured than copepods). After an encounter with cirri of feeding barnacles, copepods were usually able to swim out of the barnacles capture zone within one video frame (40 ms), by accelerating from a slow swimming speed (approximately 1.85 cm s^-1) to a mean escape swimming speed of 18.11 cm s^-1 (ca. 360 body lengths s^-1). This was not the case for A. salina nauplii, which usually remained in contact with cirri before being transferred to the mouth and ingested. Thus, experimental studies addressing the methodology of organisms feeding on zooplankton should consider that slow-swimming prey like Artemia sp. nauplii may only represent the non-evasive fraction of natural mesozooplankton assemblages.     

A novel method for investigating the collective behaviour of fish: introducing ‘Robofish’

Collective animal behaviour has attracted much attention recently, but cause-and-effect within interaction sequences has often been difficult to establish. To tackle this problem, we constructed a robotic fish (‘Robofish’) with which three-spined sticklebacks (Gasterosteus aculeatus L.) interact. Robofish is a computer-controlled replica stickleback that can be programmed to move around a tank. First, we demonstrated the functioning of the method: that the sticklebacks interacted with Robofish. We examined two types of interaction: recruitment and leadership. We found that Robofish could recruit a single fish from a refuge and could initiate a turn in singletons and in groups of ten, i.e. act as a leader. We also showed that the influence of Robofish diminished after the first 30 min that fish spent in a new environment. Second, using this method, we investigated the effects of metric and topological inter-individual distance on the influence that Robofish had on the orientation of fish in a shoal of ten. We found that inter-individual interactions during this turn were predominantly mediated by topological, rather than metric, distance. Finally, we discussed the potential of this novel method and the importance of our findings for the study of collective animal behaviour.     

Induction of swarming ofNereis succinea

Nereis succinea (Frey and Leuckart, 1847), collected in 1987 from the Weser estuary, FRG, was exposed to different temperatures in the laboratory. Metamorphosis to heteronereid stages, as well as swarming at a minimum temperature of 12°C, was induced by raising temperatures around the time of the new moon. Lunar periodicity was illustrated under natural temperature-programs, and at 16°C. An abrupt increase in temperature caused swarming to occur at different times of the lunar cycle.     

Lack of kin recognition in swarming honeybees ( Apis mellifera )

Honeybee colonies reproduce by colony fission and swarming. The primary swarm leaves the nest with the mated mother queen. Further “after-swarms” can leave the nest. These are composed of virgin queens and sister workers. Since all workers in the primary swarm have the same relationship to the mother queen, kin recognition cannot have any effect on the worker distribution in the swarm. Because of polyandry of the mother queen, the after-swarm is composed of super- and halfsister workers of the virgin queen. In this case kin recognition might affect swarm composition if workers increase their inclusive fitness by preferentially investing in a supersister queen. The distribution of workers in the mother colony, the primary and the after-swarm was analyzed using single-locus DNA fingerprinting in two colonies of the honeybee (Apis mellifera). The colonies were composed of 21 and 24 worker subfamilies because of multiple mating of the queen. The subfamily distribution in the mother colonies before swarming was significantly different from the subfamily frequencies in the primary swarm. This indicates different propensities for swarming in the various subfamilies. The subfamily distribution was also significantly different between the mother colony and the after-swarm. There was however no significant difference between the subfamily composition of the primary and the after-swarm. The average effects of kin recognition on the distribution of the subfamilies in the two after-swarms were less than 2%. We conclude that colony-level selection sets the evolutionary framework for swarming behaviour. Received: 22 May 1996 / Accepted after revision: 2 November 1996     

Experimental analysis of Froude number effect on air entrainment in the hydraulic jump

A hydraulic jump is characterized by strong energy dissipation and mixing, large-scale turbulence, air entrainment, waves, and spray. Despite recent pertinent studies, the interaction between air bubbles diffusion and momentum transfer is not completely understood. The objective of this paper is to present experimental results from new measurements performed in a rectangular horizontal flume with partially developed inflow conditions. The vertical distributions of the void fraction and the air bubbles count rate were recorded for inflow Froude number Fr ₁ in the range from 5.2 to 14.3. Rapid detrainment process was observed near the jump toe, whereas the structure of the air diffusion layer was clearly observed over longer distances. These new data were compared with previous data generally collected at lower Froude numbers. The comparison demonstrated that, at a fixed distance from the jump toe, the maximum void fraction C _max increases with the increasing Fr ₁. The vertical locations of the maximum void fraction and bubble count rate were consistent with previous studies. Finally, an empirical correlation between the upper boundary of the air diffusion layer and the distance from the impingement point was derived.     

Comparison of the food of Triphoturus mexicanus and T. nigrescens,two lanternfishes of the Pacific Ocean

Prey (chiefly euphausiids and copepods) eaten by two myctophids (lanternfishes) are compared from incidence in fish stomachs and from abundance in the environment. One lanternfish species, Triphoturus mexicanus, lives in the California Current, and the other, T. nigrescens, lives in the central Pacific Ocean. Although these two environments are very different physically and biologically, the feeding habits of the two lanternfishes are surprisingly similar. Prey biomass is 94% euphausiids, 3% copepods, and 3% other organisms for T. mexicanus and 88% euphausiids, 4.5% copepods, and 7.5% other organisms for T. nigrescens; the difference between the fish species is not significant when tested statistically. The two fishes resemble one another in frequency distributions of ingested copepod individuals, copepod species, euphausiid individuals, and euphausiid species. During a single diurnal feeding period, both fishes eat a variety of copepod species but tend to eat only a single species of euphausiid. T. mexicanus grows to twice the length of T. nigrescens and eats proportionally larger euphausiids; however, both fishes eat copepods having the same median size. The frequencies of euphausiid species in the diets of both fishes differ from the frequencies in the environment. The chief differences between the feeding habits of the two lanternfishes are that T. nigrescens, in comparison to its congener, eats a greater variety of organisms during one diurnal feeding period and captures smaller euphausiids. The feeding patterns for each lanternfish species are consistent over distances of hundreds of kilometers and over many years of sampling.     

Shadow cinematography of fish larvae

A simple system of shadow cinematography, consisting of a small tungsten halogen lamp, 2 large biconvex lenses and a 16 mm camera, is described for recording the swimming and feeding behaviour of larval fish. The system can be used either with infra-red film to record swimming behaviour independently of ambient light intensity, or with high-resolution film to record food organisms and feeding behaviour. Small plankton organisms of 0.2 mm width can be resolved using high-resolution film. The technique has been used to record the behaviour of plaice larvae (Pleuronectes platessa L.) feeding on the nauplii of Artemia salina L. The perceptive field of the larvae extends to approximately ±60° in azimuth, ±40° in elevation and 1.5 body lengths in range.     

Environmetrical interpretation of analytical data of marine organisms from the black sea

The environmetrical analysis of the benthic organisms data set from a Black sea region has revealed new information concerning the chemical content and the bioindicating abilities of polychaeta (Melina palmata), Crustacea (Aspendopsis ostroumovi) and molluscs (Mytilus gallo‐provincialis). The application of various multivariate statistical approaches like cluster and principal component analysis, linear regression and partial least square modeling, source apportioning makes it possible to understand in a better way the properties of the benthic organism as collectors of pollutants in a total and a more specific mode. It is shown that heavily polluted coastal zones are indicated in the same way by all benthic species but some specificity could be detected when moderately polluted zones are considered. In this case polychaeta accumulated preferably Co, Cr, Cu and Pb Crustacea ‐ As, Cd and Ni and molluscs ‐ Zn to a limited extent.

PCA identified three latent factors (“anthropogenic flotation”;, “anthropogenic galvanic”; and “naturally occurring") which explain about 65% of the total variance of the system and determine the data set structure. The source apportioning on the absolute principal component scores proved that none of the metals is quantitatively linked with only one anthropogenic or natural source.

The linear regression and PLS models have indicated that a reliable prognosis of the pollution on some naturally occurring chemical components (e.g. linear regression on Zn for Cr and Ni) or combination of them (PLS modeling on Mn/Zn or on Mn/Zn/Fe for the rest of the pollutants) could be achieved.     

Genetic variation among populations of Mytilus spp. in eastern Newfoundland

Small-scale population densities of tidal creek eastern mudsnails, Ilyanassa obsoleta Say (studied in 1986 and in 1992 at West Meadow Creek, Stony Brook, New York) corresponded more to variation in water flow velocity than to surface sediment chlorophyll a. Higher densities were found at low flow sites. Short-term behavioral responses are likely to be responsible for density variation. Experiments using laboratory flumes and field observations both demonstrated that the snails responded to strong flow by burrowing into the substratum. Burrowing may prevent dislodgment from the sediment surface, but it also appears to be disadvantageous since burrowed mudsnails have smaller amounts of food in their guts. Snails released in sites of periodic high flow conditions moved greater distances and were soon found near the quiet-water periphery of the creek, whereas snails released at the quiet-water periphery moved far less. It is not clear whether movement from the high flow site was through crawling or through hydrodynamic transport. Laboratory flume experiments demonstrated an active crawling movement towards areas of lower current velocity. This evidence suggests that strong bottom flow in the creek center results in a combined response of burial to avoid dislodgment and a net movement towards quiet water, which reduces exposure to the high velocity conditions of the creek center.     

Adaptation for accuracy or for precision? Diel emergence timing of the intertidal insect Pontomyia oceana (Chironomidae)

Synchronization of reproduction is obviously under strong selection, but it is not always clear whether the selection is for accuracy of timing, that is, minimal deviation from the environmental optimum, or alternatively, for precision of timing, that is, least extent of spread from the population mean. The former is usually adaptive for exploiting certain environmental conditions, whereas the fitness gain of the latter is usually associated with co-occurrence with conspecifics. We studied the intertidal midge Pontomyia oceana, which has a life cycle of about 30 or 45 days culminating in a highly synchronous swarming of the adult stage that lasts only about 1–2 h. We found that the external proximate factors, controlling their diel swarming, comprise two cues, that is, sunrise and sunset. These two cues, however, are not in existence to improve the accuracy of timing, as their diel swarming times differ widely, with respect to light condition or to tidal condition, in different evenings. Rather, the function is to improve the precision of timing. During the metabolic processes in preparation for emergence, waiting for the cues reduced variation in the population. The extent of dispersion in emergence (swarming) time in an evening is much smaller, when two cues are used compared with the inferred one-cue situations. The short adult life of the marine midges must have rendered mate-finding a stringent selective force despite their apparent high densities. The same principle, that is, using multiple cues to improve precision of timing, may be applicable to many long-lived, free-spawning species, for example, corals, where gametes can fertilize effectively only within a short time after release.     

Removal of nitrogen and phosphorus from textile wastewater by soilless culture of Lolium multiflorum

A technique of soilless culture for removal of total nitrogen (TN) and total phosphorus (TP) from textile wastewater using Lolium multiflorum was conducted in this research. The TN concentration decreased from 50.72 mg/L to 24.64–27.89 mg/L and TP decreased from 6.9 mg/L to 3.7–4.1 mg/L in the experimental tank with the size of 4.7 m x 1.2 m x 0.75 m. The results suggested that L. multiflorum could absorb a large amount of N and P elements from the wastewater. This technique of soilless culture has many advantages such as simple equipment, low cost, easy operation, low energy consumption, convenient management and flexible disposition.     

A general random walk model for the leptokurtic distribution of organism movement: Theory and application

The movement of organisms is usually leptokurtic in which some individuals move long distances while the majority remains at or near the area they are released. There has been extensive research into the origin of such leptokurtic movement, but one important aspect that has been overlooked is that the foraging behaviour of most organisms is not Brownian as assumed in most existing models. In this paper we show that such non-Brownian foraging indeed gives rise to leptokurtic distribution. We first present a general random walk model to describe the organism movement by breaking the foraging of each individual into events of active movement and inactive stationary period; its foraging behaviour is therefore fully characterized by a joint probability of how far the individual can move in each active movement and the duration it remains stationary between two consecutive movements. The spatio-temporal distribution of the organism can be described by a generalized partial differential equation, and the leptokurtic distribution is a special case when the stationary period is not exponentially distributed. Empirical observations of some organisms living in different habitats indicated that their rest time shows a power-law distribution, and we speculate that this is general for other organisms. This leads to a fractional diffusion equation with three parameters to characterize the distributions of stationary period and movement distance. A method to estimate the parameters from empirical data is given, and we apply the model to simulate the movement of two organisms living in different habitats: a stream fish (Cyprinidae: Nocomis leptocephalus) in water, and a root-feeding weevil, Sitona lepidus in the soil. Comparison of the simulations with the measured data shows close agreement. This has an important implication in ecology that the leptokurtic distribution observed at population level does not necessarily mean population heterogeneity as most existing models suggested, in which the population consists of different phenotypes; instead, a homogeneous population moving in homogeneous habitat can also lead to leptokurtic distribution.     

Habitats and tidal accessibility of the marine foods of dabbling ducks and brant in Boundary Bay,British Columbia

The role of birds in intertidal food webs is mediated by effects of tides on the habitat, distribution, and accessibility of food organisms. We studied effects of elevation, sediment characteristics, vegetation, and tidal cycles on waterfowl feeding and food organisms in Boundary Bay, British Columbia, Canada from September to March 1990–1992. Percent dry mass of esophageal contents was mainly Zostera japonica leaves (84%) for American wigeon (Anas americana Gmelin), and leaves of Z. japonica (57%) and Z. marina (41%) for brant (Branta bernicla L.). Northern pintails (A. acuta L.) and mallards (A. platyrhynchos L.) ate mostly the leaves, seeds, and rhizomes of Z. japonica (48 and 72%, respectively) and gastropods (12 and 28%). Green-winged teal (A. crecca L.) ate mainly amphipods (68%) and saltmarsh seeds (19%). The well-sorted fine sand (median 0.125 mm) to medium sand (0.25 mm), and the low organic carbon content (1.31±0.12% dry mass), did not appear to vary sufficiently along 2.1- and 2.6-km transects to affect macroinvertebrate distributions. Three habitat zones were recognized: (1) upper intertidal (sandflats); (2) mid-intertidal (largely monotypic Z. japonica); and (3) lower intertidal (mixed Z. japonica and Z. marina). These zones, although related to elevation, had greater influence on amphipod distributions than did elevation alone or detrital biomass. Biomass and numbers of most invertebrate foods of waterfowl were greatest in the eelgrass zones, with the upper elevational limit of Z. japonica occurring at about Mean Water Level (MWL). Accordingly, wigeon, pintails, and mallards fed mostly below MWL. Because of seasonal tidal cycles, daily accessibility of areas below MWL but within reach of surfacefeeding waterfowl decreased from a mean of 10.8 h in October to 9.6 h in December. In October only 10 d and <12 h available feeding time, whereas in December and January every day had <12 h available feeding time. Seasonal effects of tides on accessibility might limit sufficient feeding by some waterfowl species in intertidal areas, increasing their reliance on alternative feeding sites.