Modes of water manipulation by the lobate ctenophore Leucothea sp.

Examination of the lobate ctenophore Leucothea sp. has revealed new patterns of swimming and water manipulation in addition to the typical ctenophore mode of slow swimming with ctene plate (comb) ciliary propulsion. We distinguish between slow ctene propulsion and rapid ctene propulsion; the latter is accomplished by an increased ciliary beat that produces a coupled vortex wake, resulting in jet propulsion. The large oral lobes both capture prey and provide undulatory muscular propulsive power. The auricles exhibit distinct phasic synchrony and aphasic or sculling motions that generate small vortices in the water which facilitate prey capture. Distinctive papillae covering the exterior of Leucothea sp. may be chemo- or mechano-sensory structures. The integration of all of these structures results in an organism that is more complex behaviorally than might be expected on the basis of its superficially simple and delicate body plan. Field work involved blue-water diving in the waters of the California Bight during June and July, 1985.     

Contrast in the importance of arrow squid as prey of male New Zealand sea lions and New Zealand fur seals at The Snares,subantarctic New Zealand

New Zealand sea lions (Phocarctos hookeri) are threatened by incidental bycatch in the trawl fishery for southern arrow squid (Nototodarus sloanii). An overlap between the fishery and foraging sea lions has previously been interpreted as one piece of evidence supporting resource competition for squid. However, there is currently no consensus about the importance of squid in the diet of New Zealand sea lions. Therefore, we investigated this importance independently of spatial and temporal differences in squid availability through a simultaneous study with sympatric New Zealand fur seals (Arctocephalus forsteri), a species known to target arrow squid. Diet sampling at The Snares (48°01′S 166°32′E), subantarctic New Zealand, in February 2012 coincided with peak annual catch in the nearby squid fishery. Diets were deduced by analyses of diagnostic prey remains from scats (faeces) and casts (regurgitations). The contribution of each prey species to the diet was quantified using the per cent index of relative importance (% IRI) that combined frequency of occurrence, mass and number of prey items. Arrow squid was a minor component in sea lion scats (2 % IRI), and none was found in their casts. In contrast, arrow squid was the major component in fur seal scats and casts (93 and 99 % IRI, respectively). This study found that New Zealand sea lions ate minimal squid at a time when squid was clearly available as evidenced by the diet of New Zealand fur seals; hence, there was no indication of resource competition between sea lions and the squid fishery.     

Characteristics of residual organochlorine pesticides in soils under different land-use types on a coastal plain of the Yellow River Delta

The residual levels of organochlorine pesticides (OCPs) were examined in soils covering five types of land use along a salinity gradient on the Yellow River Delta. The most prominent OCPs were dichlorodiphenyltrichloroethane (∑DDT, arithmetic mean = 5.11 μg kg^?1), hexachlorocyclohexane (∑HCH, 1.69 μg kg^?1) and ∑endosulfan (10.4 μg kg^?1). The spatial variability of OCPs composition shifted from γ-HCH and o,p′-DDT dominated pesticides in coastal soils to p,p′-DDE dominated pesticides in inland soils. In different land-use types, the percentages of β-HCH and p,p′-DDE are characterized by more recalcitrant components in decreasing order of vegetable fields, cereal fields, cotton fields, wetlands and tidal flats with increasing soil salinity. However, the less recalcitrant components, γ-HCH and o,p′-DDT, showed an opposite trend. Endosulfan sulfate predominated in all land-use types. Residual levels of β-HCH were affected by soil organic matter. The correlations between γ-HCH and clay content and between p,p′-DDE, o,p′-DDT and salinity might associate with the influence of sediment cotransport by the Yellow River and the density of anthropogenic activities in coastal region. Depth distribution of the OCPs in typical soil profiles also implied that local historical usage and sediment transport by the Yellow River both affected the OCPs residual in this region.     

The effects of partial mortality on the fecundity of three common Caribbean corals

The recent intensification of human disturbances in the Caribbean has increased the prevalence of partial mortality on coral colonies. Partial mortality can change colony size by directly shrinking colonies or by splitting colonies into fragments. A reduction in colony size can also adversely affect fecundity and fitness as internal resources shift away from reproduction toward colony maintenance. This study aimed to determine whether three Caribbean coral species, Siderastrea siderea, Montastraea faveolata, and Diploria strigosa, along the reef tract in Puerto Morelos, Mexico (20^o52′N, 86^o51′W), continued to dedicate resources to reproduction when colonies were fragmented to pre-maturation size. Contrary to expectations, eggs were found in colonies that were smaller than the maturation size and had been subjected to partial mortality. The continued dedication of resources toward reproduction, even in the smallest colonies, suggests that resource trade-offs away from reproduction are not as rigid as previously suggested in stressed corals.     

The trophic link between squid and the emperor penguin <Emphasis Type="Italic">Aptenodytes forsteri</Emphasis> at Pointe Géologie,Antarctica

Cephalopod beaks retrieved from stomachs of dead emperor penguin chicks at Pointe Géologie, Terre Adélie, provide information on taxonomic and size composition of the penguin’s squid diet, on the trophic range of the squid species preyed upon and on the fractional trophic impact of the penguin on the whole food web. Emperor penguins prey upon four squid species (Psychroteuthis glacialis, Kondakovia longimana, Gonatus antarcticus, Alluroteuthis antarcticus) and do not take squid larger than 480 mm mantle length. Larger squid live either below the penguin’s diving range or are beyond its handling capacity. Nitrogen stable isotope ratios indicate that squids cover a range of about two trophic levels (2.5–8‰ δ¹⁵N). The impact of the emperor penguin, however, concentrates on the upper part of this range, about 68% of its squid prey being >6‰ δ¹⁵N. The principal components of the emperor’s diet, fish, krill and squid, differ distinctly in average trophic level. Consequently the trophic position of the emperor penguin changes accordingly with diet composition and may differ by almost one trophic level between different emperor penguin colonies.     

Impact of ocean warming on the early ontogeny of cephalopods: a metabolic approach

The impact of a realistic warming scenario on the metabolic physiology of early cephalopod (squid Loligo vulgaris and cuttlefish Sepia officinalis) life stages was investigated. During exposure to the warming conditions (19 °C for the western coast of Portugal in 2100), the increase in oxygen consumption rates throughout embryogenesis was much steeper in squid (28-fold increase) than in cuttlefish (11-fold increase). The elevated catabolic activity–accelerated oxygen depletion within egg capsules, which exacerbated metabolic suppression toward the end of embryogenesis. Squid late-stage embryos appear to be more impacted by warming via metabolic suppression than cuttlefish embryos. At all temperature scenarios, the transition from encapsulated embryos to planktonic paralarvae implied metabolic increments higher than 100 %. Contrary to the nektobenthic strategy of cuttlefish newborns, the planktonic squid paralarvae rely predominantly on pulsed jet locomotion that dramatically increases their energy requirements. In the future, hatchlings will require more food per unit body size and, thus, feeding intake success will be crucial, especially for squid with high metabolic rates and low levels of metabolic reserves.     

Vertical dense jet in flowing current

The discharge of brackish water, as a dense jet in a natural water body, by the osmotic power plants, undergoes complex mixing processes and has significant environmental impacts. This paper focuses on the mixing processes that develop when a dense round jet outfall perpendicularly enters a shallow flowing current. Extensive experimental measurements of both the salinity and the velocity flow fields were conducted to investigate the hydrodynamic jet behavior within the ambient current. Experiments were carried out in a closed circuit flume at the Coastal Engineering Laboratory (LIC) of the Technical University of Bari (Italy). The salinity concentration and velocity fields were analyzed, providing a more thorough knowledge about the main features of the jet behavior within the ambient flow, such as the jet penetration, spreading, dilution, terminal rise height and its impact point with the flume lower boundary. In this study, special attention is given to understand and confirm the conjecture, not yet experimentally demonstrated, of the development and orientation of the jet vortex structures. Results show that the dense jet is almost characterized by two distinct phases: a rapid ascent phase and a gradually descent phase. The measured flow velocity fields definitely confirm the formation of the counter-rotating vortices pair, within the jet cross-section, during both the ascent and descent phases. Nevertheless, the experimental results show that the counter-rotating vortices pair of both phases (ascent and descent) are of opposite rotational direction.     

Effects of dissolved ammonium addition and host feeding with Artemia salina on photoacclimation of the hermatypic coral Stylophora pistillata

 Effects of nutrient treatments on photoacclimation of the hermatypic coral Stylophora pistillata (Esper) were studied. Studies on photoacclimation of colonies from different light regimes in the field were evaluated and used to design laboratory experiments. Coral colonies were collected in the Gulf of Eilat (Israel) from January to March 1993. Exterior branches of colonies from different depths (1 to 40 m) displayed different trends in production characteristics at reduced and very low levels of illumination. From 24 ± 3% to 12 ± 2% of incident surface photosynthetic active radiation (PAR_o), zooxanthella population density and chlorophyll a+c per 10⁶ zooxanthellae increased, a trend seen in the range of light levels optimal for coral growth (90 to 30% PAR_o). The P _max of CO₂ per 10⁶ zooxanthellae decreased, while P _max of CO₂ per 10³ polyps increased, indicating an increase in zooxanthella population density at low light levels. Proliferous zooxanthella frequency (PZF, a measure of zooxanthella division) declined significantly at light levels <18 ± 3% PAR_o. At the lowest levels of illumination (<5% PAR_o), zooxanthella population density decreased, as did the PZF; chl a+c per 10⁶ zooxanthellae was unchanged. In 28-d experiments, exterior coral branches from the upper surfaces of colonies from 3 m depth (65 ± 4% PAR_o) were incubated in aquaria under bright (80 to 90% PAR_o), reduced (20 to 30% PAR_o), and extremely low (2 to 4% PAR_o) light intensities. At each light intensity, the corals were maintained in three feeding treatments: sea water (SW); ammonium enriched SW (SW + N); SW with Artemia salina nauplii (SW + A). An increase in P _max of CO₂ per 10³ polyps was found in corals acclimated to reduced light (20 to 30% PAR_o) in nutrient-enriched SW, while in SW, where the increase in zooxanthella population density was smaller, it did not occur. Nutrient enrichments (SW + N at 2 to 4% PAR_o and SW + A at 20 to 30% PAR_o) increased zooxanthella population density, but had no effect on chl a+c per 10⁶ zooxanthellae. Acclimation for 14 d to reduced (10 to 20% PAR_o) and extremely low (1 to 3% PAR_o) light intensities shifted ¹⁴C photoassimilation into glycerol and other compounds (probably glycerides), rather than sugars. Both ammonium addition and feeding with Artemia salina nauplii resulted in an increase in photosynthetic assimilation of ¹⁴C into amino acids. We conclude that acclimation to reduced light consists of two processes: an increase in photosynthetic pigments and in zooxanthella population density. Both processes require nitrogen, the increase in zooxanthella population density needing more; this adaptation is therefore limited in nitrogen-poor sea water. Received: 19 June 1998 / Accepted: 13 June 2000     

Comparative foraging ecology and ecological niche of a superabundant tropical seabird: the sooty tern Sterna fuscata in the southwest Indian Ocean

Over 6-million pairs of sooty terns Sterna fuscata breed once a year in the southwest Indian Ocean, mostly on three islands of the Mozambique Channel (Europa, Juan de Nova and Glorieuses) and in the Seychelles region. Seasonal reproduction in either winter or summer is the dominant strategy in the area, but non-seasonal reproduction also occurred in some places like at Glorieuses Archipelago. The feeding ecology of the sooty tern was investigated during the breeding seasons to determine whether terns showed significant differences in their trophic ecology between locations. Regurgitations were analyzed to describe the diet of individuals when breeding, and stable isotopes and mercury concentrations were used to temporally integrate over the medium-term of the trophic ecology of both adults and chicks. Overall, the diet was composed of fish, flying squid and fish larvae in different proportions. At Europa and Aride in the Seychelles, where winter reproduction occurs, large epipelagic prey like flying fish or squid dominated the diet. At Juan de Nova, sooty terns reproduce in summer and rely mostly on fish larvae. At Glorieuses (non-seasonal breeding), the diet was intermediate with fish larvae and flying squid being important prey items. The stable-carbon and nitrogen isotope values in blood confirm the differences observed in dietary analysis, and demonstrate different feeding strategies between colonies. δ¹³C values of feathers showed spatial segregation between birds from the Mozambique Channel and the Seychelles region. Terns from the Seychelles had also higher δ¹⁵N values. Feather δ¹³C values also suggest a significant shift from summer to wintering habitat for birds from Juan de Nova. This study emphasizes the high phenotypic plasticity of the species, which may explain its numerical dominance in all tropical waters of the World’s Ocean.     

Contrasting C and N isotope ratios from sperm whale skin and squid between the Gulf of Mexico and Gulf of California: effect of habitat

To investigate feeding variation between populations of an apex oceanic predator, stable isotope ratios of carbon (δ¹³C) and nitrogen (δ¹⁵N) have been compared in skin of female and immature male sperm whales (Physeter macrocephalus) from the northern Gulf of Mexico (GoM) and the Gulf of California (GC). Whale sexes were determined genetically. The δ¹³C and δ¹⁵N values from squid muscle were used from the GC, and from inshore and offshore sites in the GoM. We documented contrastingly lower δ¹³C and δ¹⁵N from whales and squid of the GoM compared with those from the GC. While this difference may be associated with variation in trophic position, geographic variation in biochemical cycling influenced significantly the contrasting isotope values between gulfs. Within the northern GoM, the highly distinct δ¹⁵N values of neritic squid versus mesopelagic squid provide further evidence of habitat specificity in δ¹⁵N.     

Are behavioral syndromes invariant? Spatiotemporal variation in shy/bold behavior in squid

Behavioral syndromes are correlated suites of behavior, analogous to human personality traits. Most work to date has been taken from limited “snapshots” in space and time, with the implicit assumption that a behavioral syndrome is an invariant property, fixed by evolutionary constraints or adaptations. However, directional selection on two mechanistically independent traits (selective covariance) could also result in correlated behaviors. Previously, we have shown that shy/bold behavior in Southern dumpling squid (Euprymna tasmanica) across predator encounter and feeding risk contexts is genetically and phenotypically uncoupled, and hence potentially free to vary independently. Here, we collected data on shy/bold behaviors from two independent wild populations of squid in two different years to test whether behavioral correlations across these same two functional contexts vary through time and space. We detected significant influences of population, sex, and body size on the expression of boldness in squid within each functional context, and this was coupled with significant differences in relative population density and adult sex ratio. Despite these changes in behavior and demographic parameters, we found that correlations between boldness scores across the two functional contexts were largely absent in both wild populations of squid in both years. Our work suggests that some animal groups may be largely characterized by context-specific behavioral expression. A theoretical framework which conceptualizes behavioral syndromes resulting from context-specific behavioral rules may be needed to fully understand why behaviors are sometimes correlated, and why sometimes they are not.     

Spatial and temporal variation in growth rates and maturity in the Indo-Pacific squid <Emphasis Type="Italic">Sepioteuthis lessoniana</Emphasis> (Cephalopoda: Loliginidae)

Age, growth and maturity parameters are described for the Indo-Pacific squid Sepioteuthis lessoniana both temporally and spatially (equatorial, tropical and subtropical). Tropical squid that grew through periods of warming water temperatures grew 9% faster than squid that grew through periods of cool water temperatures. The tropical spring-hatched and equatorial squid had similar growth rates (3.24, 3.18 g/day) and these were significantly faster than the tropical summer/autumn hatched squid (2.89 g/day). The oldest squid aged was 224 days, but the majority of individuals were <200 days. Subtropical squid were larger, older and matured later than equatorial and tropical spring-hatched squid. The mean weight of subtropical squid aged between 100 and 150 days was >400 g, 85% greater than tropical and equatorial squid. Geographical differences revealed that subtropical mature winter females and males had mean ages >150 days, respectively 17% and 23% older than their tropical mature winter counterparts. Temporal differences in age at maturity were also evident with tropical winter females and males having a mean age of ~140 days, respectively 41% and 25% older than their summer counterparts. Cooler subtropical and winter tropical squid had the heaviest gonads (>15 g ovaries, >1.5 g testes) compared to summer tropical and equatorial squid. However, relative gonad investment (GSI) values of the cooler squid were significantly lower with cool subtropical and winter tropical females having GSI values <3, which was about half the value of the warmer water females. This study revealed considerable plasticity in the size-at-age of this species. The tropical population had growth parameters that fluctuated between an equatorial strategy (fast growth, small body size, and small gonads) and a subtropical strategy (large body size, slower growth, and large gonads) depending on season.     

Sperm storage and mating in the deep-sea squid Taningia danae Joubin, 1931 (Oegopsida: Octopoteuthidae)

Spermatangium implantation is reported in the large oceanic squid Taningia danae, based on ten mated females from the stomachs of sperm whales. Implanted spermatangia were located in the mantle, head and neck (on both sides) or above the nuchal cartilage, under the neck collar and were often associated with incisions. These cuts ranged from 30 to 65 mm in length and were probably made by males, using the beak or arm hooks. This is the first time wounds facilitating spermatangium storage have been observed in the internal muscle layers (rather than external, as observed in some other species of squid). The implications of these observations for the mating behavior of the rarely encountered squid T. danae are discussed.     

Acute nitrate poisoning in two cattle

Abstract

The rate and product composition during photooxidation of aromatic hydrocarbons (toluene, o‐, m‐, p‐xylene and cumene) dissolved in n‐hexane and spread as a liquid film on water is reported. The photo degradability of these chemicals is 10–20%. The products identified are oxygenated aromatic compounds. It is observed that the absorption bands of the compounds under investigation depend upon the partial pressure of dissolved oxygen and these bands extend up to 290 nm. The new absorption band around 290 nm is probably due to broadening of ¹L_b bands which is responsible for direct photolysis.

stream. We have been especially interested in investigating the rate of disappearance and distribution of main oxidation products in two phases. Besides that an attempt has been made to throw some light on the reaction mechanism.     

Luminescent flashing in the midwater squids Pterygioteuthis microlampas and P. giardi

The luminescent flashing capabilities and swimming behavior were examined for two species of small midwater squids (Pterygioteuthis microlampas and P. giardi). Recently captured squid were placed in a small aquarium, where their movements and luminescent flashes could be recorded with an image intensifier, a photomultiplier tube and associated equipment. Flashing behavior was initiated by disturbing the squid with an eletrical shock. The following 4 behaviors were observed: (1) a short flash and a rapid departure from the electrode; (2) a short flash and a delayed, slow departure from the electrode; (3) a longer flash persisting after a rapid departure from the electrode; (4) a longer flash when departure was prevented by continuing the shock. Luminescent flashes probably have a general defensive function, discouraging or disrupting an attack by potential predators, and within this context there may exist numerous strategies involving flashes.     

Determination of δ13C and δ15N and trophic fractionation in jellyfish: implications for food web ecology

Application of stable isotope analysis (SIA) in jellyfish allows definition of trophic patterns not detectable using gut content analysis alone, but analytical protocols require standardization to avoid bias in interpreting isotopic data. We determined δ¹³C and δ¹⁵N in Aurelia sp. from the northern Gulf of Mexico (30°00′N, 89°00′W–30°24′N, 88°00′W) to define differences in stable isotope composition between body parts and whole body, the effect of lipid extraction on δ¹³C in tissues, and fractionation values from medusa to prey. The isotopic composition of bell and whole Aurelia sp. was not different. The increase in δ¹³C values after lipid removal suggested a correction is needed. To aid future analyses, we derived a correction equation from empirical data for jellyfish samples. Laboratory feeding experiments indicated medusae increased +4 ‰ in δ¹³C and +0.1 ‰ in δ¹⁵N compared to their diet. These results suggest protocols commonly applied for other species may be inaccurate to define Aurelia sp. trophic ecology. Because Aurelia spp. are commonly found in marine ecosystems, accurately defining their trophic role by use of SIA has implications for understanding marine food webs worldwide.     

Removal of elemental mercury by KI-impregnated clay

This study described the use of clay impregnated by KI in gas phase elemental mercury (Hg^o) removal in flue gas. The effects of KI loading, temperature, O₂, SO₂ and H₂O on Hg^o removal were investigated using a fixed bed reactor. The Hg^o removal efficiency of KI-clay with 3% KI loading could maintain at a high level (approximately 80 %) after 3 h. The KI-clay demonstrated to be a potential adsorbent for Hg^o removal when compared with activated carbon based adsorbent. O₂ was found to be an important factor in improving the Hg^o removal. O₂ was demonstrated to assist the transfer of KI to I₂ on the surface of KI-clay, which could react with Hg^o directly. NO and SO₂ could slightly improve Hg^o removal, while H₂O inhibited it greatly. The results indicated that after adsorption, most of the mercury escaped from the surface again. Some of the mercury may have been oxidized as it left the surface. The results demonstrated that the chemical reaction primarily occurred between KI and mercury on the surface of the KI-clay.     

Differential increment-deposition rate in embryonic statoliths of the loliginid squid Loligo vulgaris

 Apart from one study that reported growth of less than one increment per day in statoliths of the squid Alloteuthis subulata, most studies so far have presumed that one increment was laid down per day in the statoliths of the squid species they examined. The present study provides evidence of differential daily growth rates in embryonic statoliths of the squid Loligo vulgaris Lamarck, 1798, thus confirming a previous report for A. subulata. Incremental growth rates of L. vulgaris statoliths differ as a function of temperature. Squid embryos were incubated in the laboratory at three temperatures (12.0, 15.5 and 21.1 °C), and tetracycline staining was used to follow statolith growth. This growth slowed in squid exposed to the lowest temperature, but recovered when the squid were returned to warm conditions, indicating statolith adaptation. Statolith growth rate after incubation at 12 °C was 1.3% d⁻¹ and reached 6.1% d⁻¹ for squids exposed to 21.1 °C. Statoliths from embryos incubated at 15.5 °C yielded a rate of 1 increment d⁻¹ and a mean daily growth of 2.2 μm in the dorsal dome area of the statolith. In contrast, the slow growth of statoliths incubated at 12 °C yielded a mean daily growth of only 0.9 μm in the dorsal dome and the readings resulted in a less-than-daily increment-deposition rate. Received: 9 October 1999 / Accepted: 30 March 2000     

外源Hg胁迫对3种烟草品种叶绿素含量和光合效应的影响

为研究不同浓度汞(Hg)对烟草光合作用的影响,选择福建省3个主栽烟草品种(翠碧1号、K326、云烟87)幼苗进行盆栽试验,设置0、1、2、4、8、16 mg·kg~(-1)共6个浓度组。结果表明:(1)3种烟草叶片叶绿素含量与对照比均无显著差异(P0.05),但F_v/F_m、F_v/F_o、PI_(ABS)随胁迫浓度增加均呈下降趋势,可见Hg胁迫下烟草叶绿素含量和光合强度没有必然的相关性。(2)在较高浓度(4 mg·kg~(-1))Hg胁迫下,OJIP曲线改变了形状,表明Hg胁迫对烟草叶片光合机构影响主要是PSⅡ及光系统反应中心受到损伤,光合电子传递过程受到抑制,这可能是烟草叶片光合效应显著下降的主要因素。(3)烟草叶片PSII单位反应中心比活性参数ABS/RC、TRo/RC、DIo/RC与胁迫浓度正相关,ETo/RC和RC/CSo与胁迫浓度呈负相关,在较高浓度Hg胁迫下烟草叶片通过耗散过剩的光能和提高多余能量消耗量来保护光合机构免受更大的伤害,表现出较强的自我调节能力。     

Echolocation and foraging behavior of the lesser bulldog bat, Noctilio albiventris : preadaptations for piscivory?

We studied variability in foraging behavior of Noctilio albiventris (Chiroptera: Noctilionidae) in Costa Rica and Panamá and related it to properties of its echolocation behavior. N. albiventris searches for prey in high (>20 cm) or low (<20 cm) search flight, mostly over water. It captures insects in mid-air (aerial captures) and from the water surface (pointed dip). We once observed an individual dragging its feet through the water (directed random rake). In search flight, N. albiventris emits groups of echolocation signals (duration 10–11 ms) containing mixed signals with constant-frequency (CF) and frequency-modulated (FM) components, or pure CF signals. Sometimes, mostly over land, it produces long FM signals (duration 15–21 ms). When N. albiventris approaches prey in a pointed dip or in aerial captures, pulse duration and pulse interval are reduced, the CF component is eliminated, and a terminal phase with short FM signals (duration 2 ms) at high repetition rates (150–170 Hz) is emitted. Except for the last pulses in the terminal phase N. albiventris avoids overlap between emitted signals and echoes returning from prey. During rakes, echolocation behavior is similar to that in high search flight. We compare N. albiventris with its larger congener, N. leporinus, and discuss behavioral and morphological specializations that can be interpreted as preadaptations favoring the evolution of piscivory as seen in N. leporinus. Prominent among these specializations are the CF components of the echolocation signals which allow detection and evaluation of fluttering prey amidst clutter-echoes, high variability in foraging strategy and the associated echolocation behavior, as well as morphological specializations such as enlarged feet for capturing prey from the water surface. Received: 21 April 1997 / Accepted after revision: 12 January 1998