Riding Langmuir circulations and swimming in circles: a novel form of clustering behavior by the scyphomedusaLinuche unguiculata

Linuche unguiculata (Schwartz) seasonally forms patches in the Caribbean Sea and Indo-Pacific Ocean. Eighteen patches of medusae varying from about 500 m² to nearly 1 km² in area, were documented along the Belize barrier reef in March and April 1987, April 1988, and March and April 1990. The shape of each patch and the inter-medusa distances varied with wind velocity. At low wind speed (<4 m s^-1) patches were elliptical or circular and the individual medusae were separated by distances of 0.5 m, whereas at higher speeds windrows were evident and medusae were closer together. Windrows probably form by horizontal advection owing to convergence by Langmuir circulations. Because individual patches might exist for up to 4 mo as they drift downwind, and because winds of sufficient speed to produce Langmuir circulations do not always occur, a mechanism is necessary to maintain patch integrity during calms. In situ observations and in vitro video recording showed that the medusae swam in horizontal, near-surface, circular, clockwise trajectories. Although swimming speed was relatively high (up to 8 cm s^-1). net displacement velocity can be low (<1 cm s^-1). Thus, circular swimming probably reduces cluster breakup. Patch formation probably improves reproductive success by reducing sperm dilution.     

Estimating digestion time in gelatinous predators: a methodological comparison with the scyphomedusa Aurelia aurita

In order to quantify the trophic impact of gelatinous predators, digestion time estimates are commonly applied to counts of prey in the guts. Three primary approaches are used, the Manual-feeding, Natural-feeding and Steady-state methods; these differ in methodology and their underlying assumptions. The criteria used to define the end-point of digestion, and the resolution at which digestion progress is observed, also vary across studies. To understand the impact of such differences, we estimate digestion times of the scyphomedusa Aurelia aurita fed adult females of the copepod Acartia tonsa using these various approaches. We find ~fourfold differences which can be attributed to bias towards the slowest rates of digestion by some end-point criteria, and overestimation from low observation resolution. Artificial manipulation and the degree to which swimming and feeding behaviour are natural may also influence estimates. We provide recommendations for those quantifying digestion times of Aurelia aurita medusae and gelatinous predators.     

Diversity of swimming behavior in hydromedusae as related to feeding and utilization of space

Feeding behaviors of the following 4 species of hydromedusae are described from field and laboratory observations: Probosidactyla flavicirrata, Stomotoca atra, Phialidium gregarium and Polyorchis penicillatus. Feeding efficiency of medusae has previously been considered equivalent to fishing with a given amount (combined tentacle length) of adhesive fishing line; however, detailed observation shows that behavior of medusae greatly modifies the fishing capacity of each species. It is hypothesized that in addition to (1) tentacle number and length, the following factors strongly influence feeding efficiency: (2) tentacle posture, (3) velocity of tentacles moving through water (4) swimming pattern of medusa, (5) streamlining effects of medusa bell on water flow, (6) diameter of prey, (7) swimming pattern and velocity of prey. Each species of hydromedusa utilizes the above factors in different combinations.Mailing address     

Circatidal swimming behavior of brachyuran crab zoea larvae: implications for ebb-tide transport

Larvae of the blue crab Callinectes sapidus and fiddler crab Uca pugilator are exported from estuaries and develop on the continental shelf. Previous studies have shown that the zoea-1 larvae of some crab species use selective tidal-stream transport (STST) to migrate from estuaries to coastal areas. The STST behavior of newly hatched larvae is characterized by upward vertical migration during ebb tide followed by a descent toward the bottom during flood. The objectives of the study were (1) to determine if newly hatched zoeae of U. pugilator and C. sapidus possess endogenous tidal rhythms in vertical migration that could underlie STST, (2) to determine if the rhythms persist in the absence of estuarine chemical cues, and (3) to characterize the photoresponses of zoeae to assess the impact of light on swimming behavior and vertical distribution. Ovigerous crabs with late-stage embryos were collected from June to August 2002 and maintained under constant laboratory conditions. Following hatching, swimming activity of zoeae was monitored in darkness for 72 h. U. pugilator zoeae displayed a circatidal rhythm in swimming with peaks in activity occurring near the expected times of ebb currents in the field. Conversely, C. sapidus zoeae exhibited no clear rhythmic migration patterns. When placed in a light field that simulated the underwater angular light distribution, C. sapidus larvae displayed a weak positive phototaxis at the highest light levels tested, while U. pugilator zoeae were unresponsive. Swimming behaviors and photoresponses of both species were not significantly influenced by the presence of chemical cues associated with offshore or estuarine water. These results are consistent with predictions based on species-specific differences in spawning and the proximity of hatching areas to the mouths of estuaries. U. pugilator larvae are released within estuaries near the adult habitat. Thus, ebb-phased STST behavior by zoeae is adaptive since it enhances export. Selective pressures for a tidal migration in C. sapidus larvae are likely weaker than for U. pugilator since ovigerous females migrate seaward prior to spawning and hatching occurs near inlets and in coastal waters.     

Patterns,correlates, and paternity consequences of extraterritorial foray behavior in the field sparrow (Spizella pusilla): an automated telemetry approach

Behavioral Ecology and Sociobiology - ᅟMales of fiddler crabs (genus: Uca) construct courtship structures using mud or sand to attract mate-searching females for underground mating. A sensory...     

Local extinction of a foundation species in a hypoxic estuary: integrating individuals to ecosystem

We integrated across individual, population, community, and ecosystem levels to understand the impact of environmental stress by tracking the foundation species Mytilus edulis in the hypoxic estuary Narragansett Bay, Rhode Island, USA. Our initial surveys revealed that the mussels occurred in nine extensive (2-28 ha) dense (814-9943 individuals/m2) subtidal reefs that attracted a diverse suite of predators (sea stars, crabs, gastropods). Hypoxia occurred in the summer of 2001, and a mussel transplant experiment revealed overall reduced growth rates of individuals, and higher mortality rates among larger mussels. At the population level, large decreases in densities and cover of mussels were correlated with dissolved oxygen concentrations, leading to extinction at one site and reductions of over an order of magnitude at others. Within one year, seven of the eight remaining populations were edged to extinction, and the previously extinct population was recolonized. At the community level, a predator exclusion experiment indicated that predation was an unimportant source of mussel mortality during the hypoxic period, in part due to the emigration of sea stars, as predicted by the Consumer Stress Model. However, mussels were too intolerant to hypoxia to have a net benefit from the predation refuge. The seasonal (summer) occurrence of hypoxia allowed sea stars to return following a lag, as predicted by a stress return time model, and the resumption of predation contributed to the subsequent extinction of mussel populations. At the ecosystem level, the initial filtration rate of the mussel reefs was estimated at 134.6 x 10(6) m3/d, equivalent to filtering the volume of the bay 1.3 times during the 26-d average residence time. That function was reduced by >75% following hypoxia. The effect of hypoxia on each level of organization had consequences at others. For example, size-specific mortality and decreased growth of individuals, and reduced filtration capacity of reefs, indicated a loss of the ability of mussels to entrain planktonic productivity and potential to control future eutrophication and hypoxia. Our study quantified patterns of loss and identified pathways within an integrative framework of feedbacks, summarized in a conceptual model that is applicable to similar foundation species subjected to environmental stress.     

The utility of statoliths and bell size to elucidate age and condition of a scyphomedusa (Cassiopea sp.)

Scyphomedusae play important roles in marine ecosystems and are of economic significance. However, no reliable techniques for estimating scyphomedusa age have been documented. This study focused on the utility of Cassiopea sp. (Cnidaria: Scyphozoa) statoliths, statocysts, and body size as proxies for age of medusae. Reared medusae of known age and a manipulative experiment were used to assess the accuracy and reliability of four measures of age: number of statoliths, size (diameter) of statoliths, area of statocyst (housing statoliths), and bell diameter. Bell diameter provided the most accurate measure of age under constant conditions, but was increasingly inaccurate under varying environmental conditions. In contrast, the average number of statoliths per medusa reflected age with relatively low accuracy, but did not vary with changes in food availability and salinity. Only temperature influenced the average number of statoliths. Comparisons of bell diameter to the number of statoliths in medusae under low food availability to those fed well showed that the ratio of medusa size to the number of statoliths can be used to recognise medusae that are relatively poorly conditioned. Statoliths, therefore, provide a tool for studying both population ecology and the influence of environmental variation on medusa growth.     

Mussels flexing their muscles: a new method for quantifying bivalve behaviour

We employed a novel technique to quantify how blue mussels Mytilus edulis react to predation risk in their environment by quantifying mussel gape using a Hall sensor attached to one shell valve reacting to a magnet attached to the other. Change in gape angle per second (CHIGA) versus gape angle plots resulted in a distribution with a boundary, which defined the maximum CHIGA of a mussel at all gape angles. CHIGA boundary plots for all individual mussels were similar in form. However, the CHIGA boundary increased in extent with mussel length (maximum CHIGA for mussel valve closures for mussels 2.98 and 79.6 mm long were −1.5 and −11°s⁻¹, respectively), showing that larger mussels opened and closed most rapidly. Mussel extract added to the seawater, a factor believed to signal predation, caused mussels to close significantly faster than otherwise (P < 0.001). This approach for assessing how mussels react to their environment indicates that mussel response to predation is graded and complex and may well indicate animal-based assessments of the trade-off between effective feeding and the likelihood of predation.     

Vertical swimming in wave-induced currents as a control mechanism of intertidal migration by a sand-beach isopod

Both the vertical and horizontal distributions ofEurydice pulchra (Leach) within the water column of the surf zone were recorded throughout complete tidal cycles on a sandy beach in North Wales during the summer of 1989. Upon emerging from the sand, the isopods tended to swim up in the water column, where transport onshore would be facilitated by the wave-induced, onshore currents which laboratory wave-tank experiments have confirmed to occur near the water surface. This combination of active and passive transport to the water's edge results in high numbers of individuals in the narrow swash zone. At and just after the time of high tide, individuals swim to the water/sediment interface where, as again confirmed by wave-tank experiments, the predominant water movement is offshore. Continued swimming near the bottom during the ebb tide before reburrowing in the sand ensures transport downshore and avoidance of stranding above the characteristic level of zonation on the shore. Vertical migrations ofE. pulchra in the water column permit differential exploitation of up and downshore currents to achieve horizontal migration.     

Patterns of courtship behavior and ejaculate characteristics in male red-winged blackbirds

Sperm competition in birds is likely to have important effects on the behavior and physiology of reproduction in both sexes. For males, such competition should select for large sperm reserves and behavioral adjustment of copulation when reserves are low. We investigated both these possibilities in free-living red-winged blackbirds (Agelaius phoeniceus), a highly polygynous species with apparently strong sperm competition. We found that the recent copulatory behavior of males did not affect the propensity to copulate with a model female. Ejaculates collected from individual males at 1-h intervals showed no evidence of sperm depletion, yet repeated ejaculates collected less than 10 min apart did. Male ejaculate size was significantly larger if it was the first one of the day (i.e., after an overnight rest). The average ejaculate size was 12.5 (±12.5 SD) million sperm. Males captured during the breeding season had an average of 111.7 (±52.8) million sperm stored in their seminal glomera. Because males average a peak copulation rate of six per female per day, in one day a male might utilize all the sperm in his seminal glomera if more than two females on his territory are fertilizable. We hypothesize that polygyny and sperm competition in this species have combined to select for rapid replenishment of the seminal glomera throughout the day, in contrast to other species that have been studied. Testis size and sperm reserves of male red-winged blackbirds are intermediate between monogamous species and species with intense sperm competition. Several possible explanations for this are discussed. Received: 21 October 1997 / Accepted after revision: 15 February 1998     

Specific targeting of host individuals by a kleptoparasitic bird

Kleptoparasitism is a tactic used to acquire food opportunistically and has been shown to provide several benefits, including greater food intake rate and the acquisition of items not normally available during self-foraging. Host individuals may differ in their ability to defend themselves against kleptoparasitic attacks and therefore identifying those host individuals that are particularly vulnerable to attack could both provide energetic benefits and increase the efficiency of kleptoparasitism as a foraging strategy. Here, we show that the kleptoparasitic fork-tailed drongo (Dicrurus adsimilis) specifically targets juveniles when following groups of cooperatively breeding pied babblers (Turdoides bicolor). Drongos give alarm calls upon sighting a predator, thus providing extra predator vigilance to foraging pied babblers. However, drongos also use alarm calls to steal food items. During kleptoparasitic attacks, drongos give false alarm calls and then swoop down to steal food items dropped by alarmed babblers. Juvenile pied babblers are particularly vulnerable to attack because they (a) spend a longer period handling prey items prior to consumption and (b) respond to alarm calls primarily by immediately moving to cover, in contrast to adults who respond by looking up and visually scanning the surrounding area. Drongos attack juvenile babblers significantly more often than adults, with attacks on juveniles more likely to result in the successful procurement of a food item. This patterns of attack suggests that drongos are able to differentiate between individuals of different age when targeting pied babblers, thus increasing the efficiency of kleptoparasitism as a foraging strategy.     

Fish dispersal in fragmented landscapes: a modeling framework for quantifying the permeability of structural barriers

Dispersal is a key determinant of the spatial distribution and abundance of populations, but human-made fragmentation can create barriers that hinder dispersal and reduce population viability. This study presents a modeling framework based on dispersal kernels (modified Laplace distributions) that describe stream fish dispersal in the presence of obstacles to passage. We used mark-recapture trials to quantify summer dispersal of brook trout (Salvelinus fontinalis) in four streams crossed by a highway. The analysis identified population heterogeneity in dispersal behavior, as revealed by the presence of a dominant sedentary component (48-72% of all individuals) characterized by short mean dispersal distance (<10 m), and a secondary mobile component characterized by longer mean dispersal distance (56-1086 m). We did not detect evidence of barrier effects on dispersal through highway crossings. Simulation of various plausible scenarios indicated that detectability of barrier effects was strongly dependent on features of sampling design, such as spatial configuration of the sampling area, barrier extent, and sample size. The proposed modeling framework extends conventional dispersal kernels by incorporating structural barriers. A major strength of the approach is that ecological process (dispersal model) and sampling design (observation model) are incorporated simultaneously into the analysis. This feature can facilitate the use of prior knowledge to improve sampling efficiency of mark-recapture trials in movement studies. Model-based estimation of barrier permeability and its associated uncertainty provides a rigorous approach for quantifying the effect of barriers on stream fish dispersal and assessing population dynamics of stream fish in fragmented landscapes.     

Relationship of oxygen consumption to swimming speed in Euphausia pacifica

Swimming efficiency (the ratio of thrust power required to overcome hydrodynamic drag to net metabolic energy expenditure) was calculated for the vertically migrating euphausiid Euphausia pacifica swimming at speeds of 1–20 cm s^–1 and at temperatures of 8° and 12°C. Efficiencies ranged from 0.014 to 2.8% at 8°C and 0.009 to 1.69% at 12°C. A comparison with efficiency in fishes 2–3 orders of magnitude larger in weight (efficiency range 10–25%) indicates that locomotion in E. pacifica is far less efficient, a probable result of the organism's small size (x=33.5 mg WW) and multiple-paddle mode of propulsion. Net cost of transport of E. pacifica is three to six times the cost of a hypothetical value for sockeye salmon. Low swimming efficiencies in zooplankton such as E. pacifica are responsible for the underestimation of zooplankton swimming costs. Multiple-paddle propulsion is less efficient than the undulatory mode of fishes.     

Caveats to quantifying ecosystem services: fruit abortion blurs benefits from crop pollination.

The recent trend to place monetary values on ecosystem services has led to studies on the economic importance of pollinators for agricultural crops. Several recent studies indicate regional, long-term pollinator declines, and economic consequences have been derived from declining pollination efficiencies. However, use of pollinator services as economic incentives for conservation must consider environmental factors such as drought, pests, and diseases, which can also limit yields. Moreover, "flower excess" is a well-known reproductive strategy of plants as insurance against unpredictable, external factors that limit reproduction. With three case studies on the importance of pollination levels for amounts of harvested fruits of three tropical crops (passion fruit in Brazil, coffee in Ecuador, and cacao in Indonesia) we illustrate how reproductive strategies and environmental stress can obscure initial benefits from improved pollination. By interpreting these results with findings from evolutionary sciences, agronomy, and studies on wild-plant populations, we argue that studies on economic benefits from pollinators should include the total of ecosystem processes that (1) lead to successful pollination and (2) mobilize nutrients and improve plant quality to the extent that crop yields indeed benefit from enhanced pollinator services. Conservation incentives that use quantifications of nature's services to human welfare will benefit from approaches at the ecosystem level that take into account the broad spectrum of biological processes that limit or deliver the service.     

Relationship of oxygen consumption to swimming speed in Euphausia pacifica

Oxygen consumption rate was measured as a function of swimming velocity for the vertically migrating euphausiid Euphausia pacifica at two temperatures (8° and 12°C) and pressures (1 and 40 atm) typical of its bathymetric distribution. Increased swimming speed (x; mh^-1) required increased oxygen consumption (y; μl O₂ mg dry weight^-1 h^-1), described by the equation y = 0.012x + 0.64 at 8°C, and by y = 0.020x + 0.85 at 12° C. The current concept of low swimming costs of zooplankton, based on determinations of dead drag in copepods, is contradicted by our measurements. Temperature had a more profound effect on metabolism at higher swimming speeds (112 m h^-1; Q₁₀=2.8) than on standard metabolism (O m h^-1; Q₁₀=2.0), indicating that activity is more costly at higher temperatures. Pressure caused a small but significant (P>0.05) rise in the relationship of respiratory rate to swimming speed at both temperatures. The energy cost of vertical migration for E. pacifica was estimated by applying our data on oxygen consumption vs swimming speed to published observations on sonic scattering layer movement and the day-night distribution pattern of this species. Results indicate that the cost of a diel migration of 254 m, through a temperature change of 4 °C (8° to 12° C), would cancel any energetic benefit gained by time spent at the lower temperature typical of daytime depth. The act of vertical migration is energetically expensive; its cost should be thoroughly considered in attempts to describe the energetics of vertically migrating species.     

Shifts in foraging behavior by a Battus philenor population: field evidence for switching by individual butterflies

Summary A bivoltine east Texas population of the pipevine swallowtail butterfly (Battus philenor) exhibits a seasonal shift in the shapes of non-host leaves upon which ovipositing females land. Field mark-recapture studies and analysis of the behavior of wild females of different ages were used to distinguish between two alternative mechanisms for the shift in the populations's predominant leaf-shape search mode: seasonal differences in the outcome of learning for successively emerging naive foragers which exhibit one preference for most of their lives vs. synchronous switching by experienced foragers from one learned preference to another. Results supported each hypothesis: (1) Since wild butterflies are short-lived, the seasonal shift in searching behavior must reflect at least partly the successive emergence of naive females that learn to prefer host species with different leaf shapes. The leaf-shape preferences of older females were, in fact, stronger and less variable than those of younger individuals. About 80% of 51 marked individuals maintained stable leaf-shape search modes over recapture events, exhibiting slightly stronger preferences in later observation periods. (2) Almost 16% of marked females switched search modes across recapture events. Unmarked females sometimes switched search modes within an observation period; switching occurred only after the discovery of the host species with a leaf shape differing from that originally preferred. Switching by individual butterflies was generally more frequent at the time an adult brood shifted from one dominant search mode to another. Individual switching within an adult brood was more common in those years in which the population's shift in predominant search mode occurred during that brood.The evolution of rapid learning by naive females and conservative switching by experienced females is discussed in relation to a quantitative model of switching dynamics.     

Governance approach to China's environmental challenges: Towards a theoretical synthesis

This paper reviewed recent research in environmental governance as a response to environmental challenges at various spatial, temporal and administrative scales. It documented the shift of approach from regulation to governance, and attempted to provide a comprehensive understanding why and how the transformation occurred. It also described major factors and forces of environmental governance, and discussed research advance in environmental governance theory. Finally, this paper summarized recent research findings on environmental governance in China, and listed policy recommendation for enhancing the governance.     

To beg or to freeze: multimodal sensory integration directs behavior in a tadpole

Effective coordination of behaviors such as foraging and avoiding predators requires an assessment of cues provided by other organisms. Integrating cues from multiple sensory modalities may enhance the assessment. We studied cue integration by tadpoles of Oophaga pumilio, which live in small arboreal water pools. In this species, mothers periodically visit their tadpoles and feed them with unfertilized eggs. When mothers visit, tadpoles beg conspicuously by vibrating until fed. However, animals other than mother frogs including potential predators may visit water pools. Thus, when a visitor appears, tadpoles must use visitor cues to decide whether to beg or to remain inactive to avoid predation. To elucidate the cues that prompt these behaviors, we videotaped behavior of O. pumilio tadpoles in response to isolated and multimodal cues. Tadpoles swam more when exposed to visual or visual and chemical cues of adult O. pumilio but only exhibited begging when exposed to visual, chemical, and tactile cues together. Visual, chemical, and tactile cues from either male or female adult O. pumilio stimulated swimming and begging, but the same cues from similarly sized heterospecific frogs did not. Lastly, tadpoles exposed to a potential predator did not beg and swam less than tadpoles with no stimulus. Together, these findings suggest that O. pumilio tadpoles use multimodal cues to modulate swimming behavior accordingly in the presence of egg provisioners, predators, and other visitors and that tadpole begging is induced by multimodal cues of conspecific frogs such that tactile and perhaps chemical cues supplement visual cues.     

Forecasting industrial emissions: a monetary approach vs. a physical approach

Forecasts of industrial emissions provide a basis for impact assessment and development planning. To date, most studies have assumed that industrial emissions are simply coupled to production value at a given stage of technical progress. It has been argued that the monetary method tends to overestimate pollution loads because it is highly influenced by market prices and fails to address spatial development schemes. This article develops a land use-based environmental performance index (L-EPI) that treats the industrial land areas as a dependent variable for pollution emissions. The basic assumption of the method is that at a planning level, industrial land use change can represent the change in industrial structure and production yield. This physical metric provides a connection between the state-of-the-art and potential impacts of future development and thus avoids the intrinsic pitfalls of the industrial Gross Domestic Product-based approach. Both methods were applied to examine future industrial emissions at the planning area of Dalian Municipality, North-west China, under a development scheme provided by the urban master plan. The results suggested that the LEPI method is highly reliable and applicable for the estimation and explanation of the spatial variation associated with industrial emissions.