Flexible defense strategies: competition modifies investment in behavioral vs. morphological defenses

Competition is predicted to affect the expression of inducible defenses, but because costs of behavioral and morphological antipredator defenses differ along resource gradients, its effects on defenses may depend on the traits considered. We tested the predictions from different defense models in tadpoles of the common frog Rana temporaria, which exhibit both types of defenses. In an outdoor experiment, we exposed the tadpoles to nonlethal predators (Aeshna dragonfly larvae) and to a gradient of intraspecific competition. Morphological responses did not follow any of the expected patterns, since investment in defense was not affected by resource level. Instead, tail depth decreased in the absence of predators. Behavioral defenses followed a state-dependent model. Overall, the defense strategy of the tadpoles revealed a shift from morphological and behavioral defenses at low tadpole density to morphological defense only at high density. This difference probably reflects the different efficiency of the defenses. Hiding is an effective means of defense, but it is unsustainable when resources are scarce. Morphological responses become more important with increasing density to compensate for the increase in behavioral risk-taking. Our results indicate that competition can strongly affect reaction norms of inducible defenses and highlight the importance of integrating ecological parameters that affect the cost-benefit balance of phenotypic plasticity.     

The measurement of respiratory electron-transport-system activity in marine zooplankton

A modification of the tetrazolium reduction method (Packard, 1971) for measurement of respiratory electron transport system activity in marine zooplankton is described. A major modification is the addition of Triton X-100 to the reaction mixture to solubilize formazan produced, thus eliminating an organic extraction step and increasing the sensitivity and versatility of the method. The kinetic parameters of the assay are presented and the apparent effects of Triton X-100 are discussed. The correlation between electron-transport-system (ETS) activity and in vivo respiration (ETS/R) for Calanus pacificus has been found to be 2.02±0.29 (standard deviation). The method is presented as providing a reliable estimate of the zooplankton component of oxygen consumption in the sea.     

Population genetic consequences of larval dispersal mode and hydrography: a case study with bryozoans

Genetic analysis of the marine bryozoans Celleporella hyalina and Electra pilosa using the RAPD technique revealed population structuring corresponding to the contrasting modes of larval dispersal. Samples of C. hyalina exhibited genetic differentiation over distances as small as 10 m, concordant with the limited dispersal predicted by a simulation model, based on the short pelagic phase of the lecithotrophic larvae and the local hydrography. In contrast, E. pilosa showed high levels of genetic heterogeneity only over much larger spatial scales, commensurate with its production of comparatively long-lived planktotrophic larvae. The population differentiation observed between samples of E. pilosa, collected from sites 70 km apart, is reconcilable with coastal water currents and frontal systems that restrict the exchange of water masses between the two sites. Hydrographic conditions and discontinuous distribution of suitable substrata therefore are seen to constrain gene flow, creating opportunities for local genetic differentiation despite the high dispersal potential of pelagic larvae. Received: 9 August 2000 / Accepted: 18 November 2000     

Amphibian embryo and parental defenses and a larval predator reduce egg mortality from water mold

Water molds attack aquatic eggs worldwide and have been associated with major mortality events in some cases, but typically only in association with additional stressors. We combined field observations and laboratory experiments to study egg stage defenses against pathogenic water mold in three temperate amphibians. Spotted salamanders (Ambystoma maculatum) wrap their eggs in a protective jelly layer that prevents mold from reaching the embryos. Wood frog (Rana sylvatica) egg masses have less jelly but are laid while ponds are still cold and mold growth is slow. American toad (Bufo americanus) eggs experience the highest infection levels. They are surrounded by thin jelly and are laid when ponds have warmed and mold grows rapidly. Eggs of all three species hatched early when infected, yielding smaller and less developed hatchlings. This response was strongest in B. americanus. Precocious hatching increased vulnerability of wood frog hatchlings to invertebrate predators. Finally, despite being potential toad hatchling predators, R. sylvatica tadpoles can have a positive effect on B. americanus eggs. They eat water mold off infected toad clutches, increasing their hatching success.     

Predator-induced female behavior in the absence of male incubation feeding: an experimental study

Parental care plasticity is critical to understanding the ecological and evolutionary influence of nest predation on life history strategies. In birds, incubation imposes a trade-off between the requirements of females (i.e., food) and egg requirements (i.e., heat and protection from predators). However, studies on this topic are rare and usually restricted to species where the male feeds the incubating female, relaxing her incubation costs. Males and females can reduce their activity at the nest to avoid detection by predators. However, females could follow two alternative antipredator strategies: to delay their return to the nest to avoid attracting attention from the potential predator or to return to the nest as soon as possible to enhance nest concealment. In this study, we manipulated the perceived risk of nest predation of incubating common blackbirds (Turdus merula), a species without incubation feeding, to study female behavioral changes induced by nest predation risk. We show experimentally that female blackbirds can reduce their nest visits in the situation with higher nest predation risk. In addition, we confirm that females significantly delay their return to the nest in the presence of a nest predator, contradicting the nest concealment hypothesis. However, our results could be interpreted as a passive antipredator response (to minimize clues given to predators) or as an active antipredator response (to search for predators to expel them from their territories).     

A regularity in trace element contents of marine zooplankton species

Four zooplankton species, three Arthropoda and one Chaetognata, from the Sea of Japan were analyzed in 1984 for eight major and fifteen trace elements, mainly by instrumental neutron activation analysis. Major and trace element contents on a dry weight basis varied little within a factor of 3.7, except for Ca (a factor of 5.7) among the four species. A log-log linearity with a slope of almost-1 was observed between mean oceanic residence time calculated from the mean dissolved river-water input _R and the concentration factor with respect to mean seawater concentration (CF _SW )for each species. The products of _R and CF _SW were nearly constant within a factor of 10, except for Br and Sb over seven orders of variation of CF _SW for each zooplankton species. It demonstrates a new regularity in trace element contents of marine zooplankton species. This relationship leads to the conclusion that the concentration factors of elements for these zooplankton species with respect to elements for these zooplankton species with respect to mean dissolved river-water concentration (CF _RW )are nearly constant within a factor of 10 with average values of 2.98 to 3.43 in logarithm.     

Salinity effects on the coexistence of cryptic species: a case study on marine nematodes

The coexistence of four cryptic species of Rhabditis (Pellioditis) marina (Nematoda: Rhabditidae) at small geographical scale challenges ecological competition theory and was therefore studied in the laboratory at two different salinities, where their performance in combined cultures was compared with that in monospecies cultures. We found that three of the four cryptic species were able to coexist, but that interspecific interactions (competition and facilitation) were common. Salinity had an effect on these interactions, with a shift from contest to scramble competition. This shift may result from an increased population development of two of the four species at the lower salinity in the monospecific cultures. This experiment demonstrates that abiotic conditions may play an important role in achieving coexistence between cryptic species and can alter the interspecific interactions between them.     

Types of larval development in marine bottom invertebrates,their distribution and ecological significance: a re-evaluation

The distribution of various types of larval development among marine bottom invertebrates has been discussed on the basis of ecological evidence by Thorson (1936, 1946, 1950, 1952) and Mileikovsky (1961b, 1965). The information at hand is reviewed anew in this paper and is re-evaluated in the light of modern pertinent literature. The interrelationships between certain larval types and their distribution are not as rigid and direct as originally assumed. This can be proved even by the copy book example of the distribution of the various forms of development among species of the coastal gastropod genus Littorina. Especially among species with wide distributional areas, local populations may exhibit greater diversity in larval types than has previously been thought. Different types of larval development have now become known to exist in different populations of opisthobranch gastropods and lamellibranchs, i.e., in invertebrate groups in which such variability had been ruled out by Thorson. Variability in the type of larval development within given species — as a function of geographical, seasonal and other environmental parameters —is also more common in other marine bottom invertebrates than formerly considered. Marine bottom invertebrates are characterized not only by the 3 main different types of larval development proposed by Thorson (pelagic, direct, viviparous), but also by a fourth type: demersal (free non-pelagic) development. This fourth type occurs at all water depths and in all geographic zones of the oceans. The most important of the 4 types is pelagic (planktotrophic) development. Thorson's rule (decrease in numbers of species possessing pelagic development from the Equator towards the Poles, and from shallow-shelf waters to greater oceanic depths) is well substantiated by new data. However, one correction is necessary: pelagic development is not completely absent in the abyssal zone, as was proposed by Thorson (1950, and later), but is represented in it by at least several species belonging to various groups of invertebrates, and is also fairly common in the bathyal zone. A detailed analysis of the distributional pattern of the different types of development of marine bottom invertebrates must further take into consideration asexual reproduction with all its different modifications. Asexual reproduction in benthonic invertebrates is ecologically significant because of its common occurrence in nature; in numerous species it is also important as a biological supplement to sexual reproduction. The vast majority of species inhabiting the shallow-shelf zone and, partly, the higher levels of the slope zone of ocean areas located roughly between the polar circles, reveals development by means of planktotrophic larval stages. In the highest latitudes and on the slopes to abyssal depths—characterized by low water temperatures, scarcity of food, increasing hydrostatic pressure and other environmental peculiarities—other types of larval development prevail and, progressively, replace pelagic development with increasing latitude or depth. The distributional patterns of the various types of development among marine bottom invertebrates form one of the most important factors determining the basic distributional dynamics of the whole benthos in all oceans, both in the geological past and at the present time.Dedicated to the memory of Professor G. Thorson —founder of modern reproductive and larval ecology of marine bottom invertebrates.     

Depth regulation of larval marine decapod crustaceans: test of an hypothesis

This study tested the hypothesis that the dimensions and symmetry of the depth regulatory window of crustacean larvae are controlled by the level of light adaptation. Responses of first and last zoeal stages of the crab Rhithropanopeus harrisii (Gould) to different rates of pressure change were analyzed with a video system. Crabs were collected from the Neuse River estuary (North Carolina, USA) from May to September 1988. Responses were measured when larvae were adapted to light having an angular light distribution similar to that underwater at intensities ranging from one log unit above the lower phototaxis threshold to four log units higher. For both zoeal stages in darkness and at 10^-6 W m^-2, the distance larvae descend before responding to a pressure increase was much shorter than the distance they would ascend before responding to a pressure decrease. When adapted to a light level of 10^-4 W m^-2 both zoeal stages descended and ascended approximately equal distances before responding to an increase or decrease in pressure, respectively. Finally at the highest test light intensity (10^-2 W m^-2), the ascent distance was much shorter than the descent distance. These results support the hypothesis. The depth regulatory window dimensions predict an ascent in the water column upon adaptation to low light intensities and descent at high light levels. Thus Sulkin's negative feedback model provides the general mechanism of depth regulation. The effects of light adaptation on the limits of the depth regulatory window provide an additional component that negates the requirement for depth regulation at an absolute depth. The composite model can be termed the light-dependent negative feedback model of depth regulation.     

Use of biomarkers in zooplankton for assessment of the 'health status' of marine and brackish environments: a short overview

The biomarker approach has been used for 25 years to study the environmental quality of marine, brackish and freshwater ecosystems. Biomarkers may indicate health status and can be applied to organisms of all zoological phyla by destructive or non destructive methods. For 5 years we have been using this approach in zooplankton to detect ecotoxicological alterations at low levels of the food chain due to contaminants. Here we review our approach to validate and apply biomarker techniques in zooplankton. We discuss advantages, limitations, some results and future research. We indicate that biomarkers in zooplankton can be used as new indices of trophic status and ecological integrity of Italian marine coastal and lagoon environments, to be included among the tools specified by Italian law D.Lgs. 152/2006.     

Combining DNA and morphological analyses of faecal samples improves insight into trophic interactions: a case study using a generalist predator

The diet of pinnipeds is most commonly inferred from morphologically diagnostic remains of prey in their scats. Although this method can generate quantitative estimates of diet simply, important prey types may not always be detected. DNA-based methods improve detection of prey in scats, but they are not quantitative. While some studies have combined morphological and DNA-based methods, these have only assessed prey that are represented by their hard remains in scats. To overcome this bias, we apply molecular and morphological analyses to the soft and hard portions of faecal samples respectively, to estimate the diet of lactating Antarctic fur seals (Arctocephalus gazella) on Heard Island. The diet of this population is of particular interest because it is expanding rapidly and may rely to some extent on mackerel icefish (Champsocephalus gunnari), which are subject to commercial fisheries. Based on results from morphological analysis and likely important prey types, we tested for DNA remains of C. gunnari, myctophids and squid in faecal samples. The proportion of samples (n = 54) yielding no dietary information was reduced from around 25.9% using either method alone, to 9.3% when combined. Detection of all prey types tested for was notably improved by integrating molecular and morphological data. Data from either method alone would have underestimated the number of animals consuming C. gunnari by around 25.7%. Detection of multiple prey types in samples increased from 9.3% when using morphological analysis only, to 33.3% when using DNA only, to 46.3% when using both methods. Taken in isolation, morphological data inferred that individual seals consume either C. gunnari or myctophids, probably foraging in separate locations characteristic of those prey. Including molecular data demonstrated that while this may be true of some individuals, many other seals consume a mixed diet of at least C. gunnari, myctophids and squid. This new approach of combining DNA-based and morphological analyses of diet samples markedly increased the number of samples yielding dietary information, as well as increasing the amount of information attained from those samples. Our findings illustrate the broad potential of this technique to improve insight into trophic interactions in marine ecosystems.

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Discriminating zooplankton assemblages through multivariate methods: A case for a tropical polluted harbour and Bar-built estuary

A comparative study (June-July 2001) on zooplankton community structure amid polluted conditions in a stagnant harbour and relatively unaffected tidal estuary near Visakhapatnam, on the east coast of India, revealed a marked disparity in species composition and abundance. While the harbour supported a rich population of calanoids (46.4%), the estuary sustained mostly cyclopoids (55.2%). Univariate and multivariate techniques (species diversity, clustering, non-metric multi-dimensional scaling and one-way ANOSIM) revealed the existence of two differing zooplankton assemblages and associated water quality (similarity 50.6%). While the estuary is typified by high amounts of dissolved silica (67.4 ± 17.7 μmol l^-1) linked with monsoon influx, the harbour waters revealed abnormal levels of phosphate (40.9 ± 9.2 μmol l^-1) and nitrate (15.3 ± 5.41 μmol l^{- 1}) suggestive of intense eutrophication, caused by the discharge of fertilizer-factory waste and domestic sewage. On the basis of routines (e.g. BVSTEP, SIMPER) implemented in Plymouth Routines in Multivariate Ecological Research, it was possible to demonstrate that while species such as Oithona rigida, Oithona brevicornis, crustacean nauplii, gastropod veligers, Acartia spinicauda, and Acartia centrura played a key role in discriminating the zooplankton assemblage in the estuary, Acrocalanus spp. (mainly Acrocalanus gracilis) played a keyed role in harbour waters. Canonical Correspondence Analysis revealed species-environment relationships; for example, while the distribution of Oithona spp. and its associates in the estuary corresponded intimately with high silicate, temperature, and low salinity, it was high salinity, phosphate, and nitrate in the harbour channel that supported a different assemblage of copepods dominated by calanoids.     

Stable isotope and biochemical fractionation in the marine pelagic food chain: the jellyfish Pelagia noctiluca and net zooplankton

In our field study we analyzed the C and H isotopic and biochemical (C, N, P, protein, lipid, carbohydrate) composition of the jellyfish Pelagia noctiluca (collected from the Gulf of Trieste in 1985 to 1986) and its presumed diet-net zooplankton. The mean ¹³C (-18.8) and D (-58.4) ratios of P. noctiluca showed enrichment in heavy isotopes relative to net zooplankton (2 for carbon and 30 for hydrogen). Both the jellyfish and net zooplankton were characterized by a linear correlation between ¹³C and D. C. N, P, protein, lipid, and carbohydrate contents of P. noctiluca were low on a dry weight basis as compared to net zooplankton. Significantly lower C:N and C:P ratios were found in jellyfish indicating a greater loss of carbon relative to nitrogen and phosphorus along the passage to a higher trophic level. Isotopic and biochemical evidence indicate that, though collected in nearshore waters, P. noctiluca depended on autochthonous marine organic matter.     

Morphological study of marine bacterial capsules: implications for marine aggregates

The morphological variation in bacterial capsules found in suspended and sinking particles from the open ocean is examined. The capsules consist of matrices of mucopolysaccharides that remain attached to and extend the cell surface. Transmission electron microscope images of several distinct and recurring morphs are presented. Capsule abundance relative to total numbers of bacteria is significantly higher in sinking particles (up to 80% of total bacteria present) than in fine suspended particles (up to 20%). Discrete fecal pellets and minipellets are inadequate sources of the capsules captured by sediment traps. Alternatively, the data and previous observations indicate that the bacterial capsules are removed from the fine suspended-particle phase in association with macroaggregates, or marine snow. The combined properties of the bacterial capsules with their extensive open matrices of attached sticky polymers suggest a high aggregation efficiency.     

Amino acid metabolism and protein turnover in larval turbot (Scophthalmus maximus) fed natural zooplankton or Artemia

The present paper studied the influence of different food regimes on the free amino acid (FAA) pool, the rate of protein turnover, the flux of amino acids, and their relation to growth of larval turbot (Scophthalmus maximus L.) from first feeding until metamorphosis. The amino acid profile of protein was stable during the larval period although some small, but significant, differences were found. Turbot larvae had proteins which were rich in leucine and aspartate, and poor in glutamate, suggesting a high leucine requirement. The profile of the FAA pool was highly variable and quite different from the amino acid profile in protein. The proportion of essential FAA decreased with development. High contents of free tyrosine and phenylalanine were found on Day 3, while free taurine was present at high levels throughout the experimental period. Larval growth rates were positively correlated with taurine levels, suggesting a dietary dependency for taurine and/or sulphur amino acids. Reduced growth rates in Artemia-fed larvae were associated with lower levels of free methionine, indicating that this diet is deficient in methionine for turbot larvae. Leucine might also be limiting turbot growth as the different diet organisms had lower levels of this amino acid in the free pool than was found in the larval protein. A previously presented model was used to describe the flux of amino acids in growing turbot larvae. The FAA pool was found to be small and variable. It was estimated that the daily dietary amino acid intake might be up to ten times the larval FAA pool. In addition, protein synthesis and protein degradation might daily remove and return, respectively, the equivalent of up to 20 and 10 times the size of the FAA pool. In an early phase (Day 11) high growth rates were associated with a relatively low protein turnover, while at a later stage (Day 17), a much higher turnover was observed. Received: 19 March 1997 / Accepted: 14 April 1997     

Physiological tolerances across latitudes: thermal sensitivity of larval marine snails (Nucella spp.)

A critical step in understanding how temperature will affect biodiversity in coastal ecosystems is to gain insight into how the tolerances, and ultimately survival, of early life history stages will influence the distribution and abundance of adults. We assessed the thermal tolerance of encapsulated veliger-stage larvae of a common dogwhelk, Nucella ostrina, that occur in the rocky intertidal zone on the west coast of North America. Results showed that veligers collected from northern latitudes in Washington State were less tolerant of heat stress than those from central sites in California. For all sites, we found there to be a subtle difference between the temperatures at which veligers first began to die compared to when veligers reached 100% mortality. On a biogeographic scale, the LT₅₀ temperatures, a measure of larval sensitivity, for N. ostrina veligers displayed a strong latitudinal trend. These findings provide a conservative measurement of the upper thermal limits of encapsulated veligers while illustrating how these early life history stages could be physiologically compromised under future climate warming scenarios.     

Summer zooplankton distribution in a Georgia Estuary

Zooplankton were collected with Van Dorn bottles at two estuarine stations near Savannah, Georgia, USA, during summer, 1974, from the surface, 2 m, 4 m, and at the bottom of the water column at 2-h intervals for 11 days. Acartia tonsa occurred at the surface at night and during high and rising tides in higher concentrations than at the same depth during daylight and other tide stages. The highest bottom concentrations of this calanoid copepod occurred at high tide. A negative phototaxic response was also apparent in this species. While not quantitatively sampled by Van Dorn bottles, harpacticoid copepods and Pseudodiaptomus coronatus Were distributed in a manner similar to that of A. tonsa. Copepod nauplii, on the other hand, were always more abundant at the surface than at the bottom, with the highest concentration occurring during low tide. The distribution pattern of nauplii appears to be a function of dilution and concentration by tidally induced changes in water volume, whereas more advanced copepod stages follow a complexly controlled distribution pattern linked with tidal mixing dynamics as outlined in a previous study by Jacobs (1968).     

A policy alternative for lagoon management: case study of Sre Ambel coastal marine fisheries,Cambodia

Sre Ambel is the most productive coastal area for marine fisheries in Cambodia. This paper explores the management concept of extractive reserve instruments to find the most effective application in the coastal areas of Cambodia. Since there are various extractive models in the world, the most important objective is to determine which extractive reserve model is best suited to real practices where the ability of the fisheries’ residents to apply laws and guidelines vary from one place to another. To reflect these objectives, this paper looks at various examples around the world, especially from various Latin American countries. The level of local ability to apply laws and regulations is determined by the weight average index method. These results shows that fishers’ education levels are sufficient to apply laws and regulations, but their understanding of existing fisheries guidelines is still below average. Although the level to which they satisfy the guidelines is poor, local people are willing to establish new appropriate guidelines for future effective application. Therefore, local fishers have the ability to apply laws and regulations under new extractive reserve aspects. An extractive reserve policy combines all available instruments to manage the lagoon. The best alternative is to develop strong collaboration among the various stakeholders in the area. This converts an open-access to common property resource use regime. All instruments of extractive reserve were satisfied by the local fishers. However, extractive reserve seems to be similar with fish refuge concept which currently proposed by Southeast Asian Fisheries Development Centre (SEAFDEC).     

Antipredator defenses along a latitudinal gradient in Rana temporaria

Antipredator defenses are expected to decrease toward higher latitudes because predation rates are predicted to decrease with latitude. However, latitudinal variation in predator avoidance and defense mechanisms has seldom been studied. We studied tadpole antipredator defenses in seven Rana temporaria populations collected along a 1500-km latitudinal gradient across Sweden, along which previous studies have found increasing tadpole growth and development rates. In a laboratory common garden experiment, we measured behavioral and morphological defenses by raising tadpoles in the presence and absence of a predator (Aeshna dragonfly larva) in two temperature treatments. We also estimated tadpole survival in the presence of free-ranging predators and compared predator densities between R. temporaria breeding ponds situated at low and high latitudes. Activity and foraging were generally positively correlated with latitude in the common garden experiment. While all populations responded to predator presence by decreasing activity and foraging, high-latitude populations maintained higher activity levels in the presence of the predator. All populations exhibited defensive morphology in body and tail shape. However, whereas tail depth tended to increase with latitude in the presence of predator, it did not change with latitude in the absence of the predator. Predator presence generally increased larval period and decreased growth rate. In the southern populations, predator presence tended to have a negative effect on metamorphic size, whereas in the northern populations predators had little or a positive effect on size. Latitude of origin had a strong effect on survival in the presence of a free-ranging predator, with high-latitude tadpoles experiencing higher mortality than those from the low latitudes. In the wild, predator densities were significantly lower in high-latitude than in mid-latitude breeding ponds. Although the higher activity level in the northern populations seems to confer a significant survival disadvantage under predation risk, it is probably needed to maintain the high growth and development rates. However, the occurrence of R. temporaria at high latitudes may be facilitated by the lower predator densities in the north.