Diurnal dynamics of the microbial loop in peatlands: structure, function and relationship to environmental parameters

Globally, introductions of alien species are increasingly common, with invasive predators potentially having detrimental effects via predation on native species. However, native prey may avoid predation by adopting new behaviors. To determine whether invasive fish populations consume endemic shrimp in invaded Hawaiian anchialine habitats or if adopted patterns of diel migration prevents predation as previously hypothesized, a total of 183 invasive poeciliids (158 Gambusia affinis and 25 Poecilia reticulata) were collected for gut content analyses from four anchialine sites during wet and dry seasons on the islands of Hawai‘i and Maui. Predation on shrimp was not detected in habitats where they retreat exclusively into the underlying aquifer diurnally and only emerge nocturnally. However, low levels of predation were detected (7/65 fishes, only by Gambusia affinis) at Waianapanapa Cave, Maui, where shrimp retreat into both the aquifer and a cave during the day. Thus, adopted behavioral responses to invasive fishes generally, though not universally, prevent predation on endemic Hawaiian anchialine shrimps. However, non-consumptive effects resulting from behavioral modification of shrimps may have appreciable impacts on the Hawaiian anchialine ecosystem and warrant further study.     

Diel variation in predator abundance, predation risk and prey distribution in shallow-water estuarine habitats

Predation by visual predators is often affected by light conditions and may therefore exhibit strong diel variation. The dominant predators on grass shrimp, Palaemonetes pugio, are finfish predators that are thought to locate their prey by visual cues. We examined the response of grass shrimp to diel variation in predation risk in the nearshore shallow waters of the Chesapeake Bay. We used diel shoreline seines to assess the relative abundance of predators. We assessed the relative risk of predation with shrimp tethered at refuge (30 cm) and nonrefuge (60 cm) depths. To measure grass shrimp response to predation risk, we used dipnets to monitor habitat use. Four predominantly visual predators dominated the shoreline seine catches, Fundulus heteroclitus, Micropogonias undulatus, Morone americana and Morone saxatilis. Total predator abundance had a diel component, with dramatic nighttime decreases in total abundance, whereas guild composition and relative abundance remained unchanged. Relative predation risk for tethered shrimp exhibited significant time by habitat interaction. During the day, depth negatively affected survivorship of tethered shrimp while at night overall survivorship increased and there was no effect of depth. Shrimp habitats use reflected diel predation risks. Abundances in the near shore were highest during the day with decreased abundances at night. Together, the seine and tethering data highlight the importance for a refuge (e.g., shallow water) from predation during the daytime and a relaxation of predation pressure at night.     

Density‐ and trait‐mediated top–down effects modify bottom–up control of a highly endemic tropical aquatic food web

Benthic invertebrates mediate bottom–up and top–down influences in aquatic food webs, and changes in the abundance or traits of invertebrates can alter the strength of top–down effects. Studies assessing the role of invertebrate abundance and behavior as controls on food web structure are rare at the whole ecosystem scale. Here we use a comparative approach to investigate bottom–up and top–down influences on whole anchialine pond ecosystems in coastal Hawai‘i. In these ponds, a single species of endemic atyid shrimp (Halocaridina rubra) is believed to structure epilithon communities. Many Hawaiian anchialine ponds and their endemic fauna, however, have been greatly altered by bottom–up (increased nutrient enrichment) and top–down (introduced fish predators) disturbances from human development. We present the results of a survey of dissolved nutrient concentrations, epilithon biomass and composition, and H. rubra abundance and behavior in anchialine ponds with and without invasive predatory fish along a nutrient concentration gradient on the North Kona coast of Hawai‘i. We use linear models to assess 1) the effects of nutrient loading and fish introductions on pond food web structure and 2) the role of shrimp density and behavior in effecting that change. We find evidence for bottom–up food web control, in that nutrients were associated with increased epilithon biomass, autotrophy and nutrient content as well as increased abundance and size of H. rubra. We also find evidence for top–down control, as ponds with invasive predatory fish had higher epilithon biomass, productivity, and nutrient content. Top–down effects were transmitted by both altered H. rubra abundance, which changed the biomass of epilithon, and H. rubra behavior, which changed the composition of the epilithon. Our study extends experimental findings on bottom–up and top–down control to the whole ecosystem scale and finds evidence for qualitatively different effects of trait‐ and density‐mediated change in top–down influences.     

Diel vertical migration of the copepod <Emphasis Type="Italic">Thermocyclops inversus</Emphasis> (Kiefer, 1936) in a tropical reservoir: the role of oxygen and the spatial overlap with <Emphasis Type="Italic">Chaoborus</Emphasis>

The effect of water transparency, dissolved oxygen concentration and the invertebrate predator Chaoborus brasiliensis on the day–night vertical distribution of the copepod cyclopoid Thermocyclops inversus was investigated in a shallow tropical reservoir, Nado Reservoir, Belo Horizonte, Brazil. Diel cycles were carried out over a period of 12 consecutive months, between October 1999 and September 2000. The different developmental stages of T. inversus exhibited diel vertical migration (DVM) and displayed a clear ontogenetic trend, with the amplitude of DVM increasing with the age of the organism, and ranging from 0.4 m to 0.8 m for nauplii, 0.4 m to 1.2 m for copepodite, and 1.1 m to 2.1 m for adults. We observed that seasonal changes in dissolved oxygen and C. brasiliensis directly influenced the vertical distribution of the copepod population in this reservoir. Furthermore, it was showed that the diurnal vertical migration is an important predator avoidance behavior since it diminished the spatial overlap between prey and its potential predator. This finding supports the hypothesis that the vertical migration is a defense mechanism against predation. Thus, T. inversus is able to remain in the anoxic layers during day light hours, and at night they move upwards avoiding hypolimnetic waters to escape from predation by Chaoborus.     

Vertical, lateral and longitudinal movement of zooplankton in a large river

1. The spatial distribution and movement patterns of zooplankton in large rivers are little known compared with those in lake environments. We conducted a series of studies in the Ohio River (U.S.A.) during the low flow period to assess diel vertical (DVM), longitudinal and lateral movement of crustacean zooplankton. 2. The dominant large zooplankter, the copepod Eurytemora affinis, showed a consistent vertical migration pattern of daytime ascent and night‐time descent during all sampling periods – the reverse of the most common migratory pattern of zooplankton in lakes. The cladoceran Bosmina migrated in a similar way in two of the three sampling periods. Surveys taken longitudinally in the river showed similar trends for both taxa. 3. During the lateral surveys, E. affinis was significantly more abundant in the shallow littoral zone during the night than in the daytime. The combination of vertical and lateral movement patterns along with the diel distribution of zooplanktivorous fish suggest that these movements are a predator‐avoidance mechanism. 4. Sampling programmes in large rivers should consider that larger zooplankton such as E. affinis may not be randomly distributed in the river channel and behaviours such as diel vertical migration may be just as evident in river habitats as in lakes.     

Diel vertical migrations of zooplankton in a shallow, fishless pond: a possible avoidance-response cascade induced by notonectids

1. Day (noon) and night (midnight) vertical distributions of zooplankton and phytoplankton in the water column (1.5 m) of a Vermont pond were determined on two consecutive days from 470 mL water samples taken at three depths (0.1, 0.5 and 1.0 m) at three sites. There was little variation across depths in temperature, dissolved oxygen concentration and phytoplankton. All individuals of each zooplankton species (a small copepod, Tropocyclops extensus and six rotifers) were counted. 2. A three-way ANOVA on the zooplankton data showed no effect of date or time of day on the abundance of any species. Significant diel shifts in vertical distribution (depth × time-of-day interactions) were found for T. extensus (nauplii, as well as copepodites and adults) and Polyarthra remata, but not for Hexarthra mira, Keratella cochlearis, Anuraeopsis fissa, Ascomorpha ovalis and Plationus patulus. Tropocyclops extensus showed a pronounced, typical diel vertical migration, avoiding the surface and occurring most abundantly near the bottom during the day. Polyarthra remata showed an equally pronounced, reverse diel vertical migration, avoiding the bottom and being most abundant near the surface during the day. 3. The diurnal descent of Tropocyclops is interpreted as an avoidance response to Buenoa macrotibialis, a notonectid which feeds on this copepod at the surface during the day but not at night. The diurnal ascent of Polyarthra is thought to be an avoidance response to Tropocyclops, which strongly suppresses this rotifer in field enclosures and laboratory vessels. Thus, these out-of-phase migrations may be coupled and represent a behavioural cascade initiated by Buenoa. 4. At night, Tropocyclops and Polyarthra both were uniformly distributed across depths. This is believed to reflect the absence of appreciable depth-related variation in temperature, algal food resources (biovolume of cryptomonads and chrysophyte flagellates) and predation risk at this time. 5. The five rotifer species that did not exhibit diel vertical migrations may be less susceptible to Tropocyclops predation than Polyarthra.     

Salinity tolerance of the gemmules of Eunapius fragilis (Leidy) and the inhibition of germination by various salts

We report results of a field test of the predator avoidance hypothesis as an explanation of the adaptive significance of diel vertical migration in zooplankton. We determined the vertical distribution and diel migration of the planktonic copepod Acartia hudsonica, concurrently with the abundance of pelagic fish, transparency and thermal stratification of the water column, on six cruises over a one year period in a temperate marine lagoon (Jakles Lagoon, San Juan Island, Washington, USA). Striking seasonal variability was observed in all biological and environmental variables. Linear regressions of the strength of diel vertical migration in A. hudsonica on these environmental variables resulted in only one statistically significant relationship, that between copepod diel vertical migration and predator abundance. These results, together with those of previous studies, point to diel vertical migration as a widespread behavioral response of planktonic prey to the presence of their predators.     

Effects of large Saduria entomon (Isopoda) on spatial distribution of their small S. entomon and Monoporeia affinis (Amphipoda) prey

We performed laboratory experiments to investigate the effects of predator avoidance and numerical effects of predation on spatial distribution of small Saduria entomon (Isopoda) and Monoporeia affinis (Amphipoda), with large S. entomon as predators. The horizontal distribution and mortality of the prey species, separately and together, were studied in aquaria with a spatial horizontal refuge. We also estimated effects of refuge on mortality of small S. entomon and M. affinis by experiments without the refuge net. In addition, we investigated whether predation risk from large S. entomon influenced the swimming activity of M. affinis, to clarify the mechanisms behind the spatial distribution. Both small S. entomon and M. affinis avoided large S. entomon. The avoidance behaviour of M. fffinis contributed about 10 times more to the high proportion in the refuge than numerical effects of predation. Due to the low mortality of small S. entomon the avoidance behaviour of this species was even more important for the spatial distribution. The combined effect of avoidance behaviour and predation in both species was aggregation, producting a positive correlation between the species in density. M. affinis showed two types of avoidance behaviour. In the activity experiments they reduced activity by 36% and buried themselves in the sediment. In the refuge experiments we also observed avoidance behaviour with the emigration rate from the predator compartment being twice the immigration rate. The refuge did not lower predation mortality in M. affinis, probably due to the small scale of the experimental units in relation to the mobility of the species. Predation mortality in small S. entomon was higher in absence of a refuge and especially high in absence of M. affinis.     

Horizontal distribution of pelagic zooplankton in relation to predation gradients

An investigation was conducted examining the horizontal and vertical distribution of zooplankton in Lake Miramar, a southern California reservoir. Daphnia and Mesocyclops populations were most abundant offshore and in deeper water during the day but appeared to move toward shore and upward at night. The results of inshore zooplankton sampling provided no evidence chat the diel horizontal migration pattern was a result of sampler avoidance by zooplankton. Inshore-offshore differences in Daphnia and Mesocyclops abundance and diel migrations were reduced during winter and early spring. Rotifer zooplankters exhibited less seasonal variation in their horizontal distributions than did the large crustacean zooplankters at all times of the year. It is hypothesized that the spatial distribution of zooplankton is related to predation gradients in Lake Miramar. The dominant planktivore in the reservoir, young-of-the-year Micropterus salmoides. was abundant from late May through December and much less so from January to early May. They were largely restricted to the littoral zone and this produced horizontal gradients of planktivory which varied in strength seasonally and from day lo night. It appears that crustacean zooplankton in Lake Miramar avoid areas with abundant planktivores during the day but migrate into these areas at night when the intensity of planktivory is reduced. Rotifers exhibit less horizontal heterogeneity and no significant diel migrations, which is attributed to the reduced risk of predation that rotifers experience relative to crustacean zooplankters. A graphical model is proposed to integrate our understanding of diel vertical and horizontal migrations of zooplankton. In this model, gradients of predation are completely vertical in offshore areas and strongly horizontal in near shore areas. Gradients of food availability are roughly similar to those of predation intensity. Plankiers respond to these gradients by migrating in a path parallel to gradients of predation at dawn and parallel to gradients of food availability after dark.     

An Unique Anchialine Pool in the Hawaiian Islands

The Sailor's Hat crater was artificially formed on the south coast of Kaho'olawe Island in 1965 with explosives. The explosion formed a crater about 50 m from the shoreline, which penetrates the watertable to a 5 m depth. The pool at the bottom of the crater meets the criteria of an anchialine pond because it shows tidal fluctuation, has measurable salinity, and lacks surface connections to the sea. The water chemistry of this pool is similar to the ocean except silica is elevated and salinity is slightly depressed suggesting a small groundwater influence. The fauna is dominated by waterboatmen, an endemic shrimp and tubeworm, polychaetes, amphipods, an ostracod, gastropod, solitary ectoproct. anemone, flatworm and sponge. The atyid shrimp, Halocaridina rubra, is a characteristic species of Hawaiian anchialine systems and probably colonized this 32-year old pool by active migration via the watertable. Colonization by the remaining fauna may have occurred by storm surf (for marine species) or with the wind. Most predators are unable to inhabit anchialine ponds because of difficult access due to physical barriers, or to unsuitable ecological conditions. The anchialine habitat and life history strategy of the atyid shrimp have probably been important influences on the adaptative success of H. rubra in the Hawaiian Islands, and may be important characteristics of hypogeal anchialine species elsewhere.     

Variation in horizontal zooplankton abundance in mountain lakes: shore avoidance or fish predation?

Hypothesizing that fish predation, active shore avoidance and outlet stream avoidance may be separately affecting horizontal zooplankton distribution, the effects of fish presence, sampling location (midlake, outlet and non-outlet shore) and time (day or night) on zooplankton abundance and body size were tested. Statistically significant horizontal zooplankton abundance gradients occurred in both fish-present and fish-absent lakes. Fish may strengthen zooplankton spatial patterns common to both fish-present and fish-absent lakes, as abundance differences among locations were often greater in fish-present systems compared to fish-absent systems. Horizontal zooplankton abundance gradients differed through a diel cycle, but were species specific with some species exhibiting gradients only during the day, while others exhibit gradients only during the night. Avoidance of the outlet over and above active shore avoidance appeared to take place in Daphnia sp. Other taxa provided equivocal support of active outlet avoidance with most showing no significant difference between shore and outlet abundance (seven of nine), one taxa showing a significant decrease and one a significant increase in outlet compared to shore abundance. No gradients in zooplankton body size were found.      

To migrate, or not to migrate: partial diel horizontal migration of fish in a temperate freshwater reservoir

The diel horizontal migration (DHM) of fish between the inshore and offshore zones of the ?ímov Reservoir (Czech Republic, deep, stratified, meso-eutrophic) was investigated by a combination of horizontal and vertical hydroacoustic surveys at 3-h intervals over 48 h and day/night purse seining in August 2007. An overwhelming majority of fish were aggregated within the epilimnetic layer. Considering only the horizontal surveys, cyclic diel fish movements between inshore and offshore habitats were apparent, while the total fish biomass remained constant between day and night. A higher fish biomass was detected in the offshore zone during daytime by both hydroacoustics and purse seining. In contrast, a higher fish biomass was recorded at night in the inshore zone. Bream Abramis brama, roach Rutilus rutilus, and perch Perca fluviatilis dominated the daytime offshore fish assemblage whereas bleak Alburnus alburnus prevailed at night. Bream and roach decreased in abundance at night while perch completely disappeared from the offshore habitat. The diel differences in size distributions and direct catches suggested the population-wide horizontal offshore migration of bleak and inshore migration of all perch during dusk. On the other hand, partial inshore migration of bream and roach adults was observed during dusk (52 and 80%, respectively). The different proportions of offshore residents among species and size classes suggested that differences in size, and, therefore, predation vulnerability, contributed to the observed migration patterns.     

Diel vertical migration in zooplankton: rapid individual response to predators

While diel vertical migration in zooplankton has been shownrecently to be a predator avoidance behavior, the mechanismby which predators induce and maintain such behavior has beendebated. We report results of an in situ predator manipulationexperiment during which enclosed populations of the marine planktomccopepod Acaraa hudsonica rapidly changed their vertical distributionand diel migration behavior depending on presence or absenceof the planktivorous fish Casterosteus aculeatus These resultspoint unambiguously to phenotypic behavioral plasticity of individualplanktonic prey, not, as previously hypothesized, population-geneticlevel behavioral changes caused by selective fish predation,as the mechanism underlying changes in diel vertical migrationin this copepod.     

Deep impact: in situ functional responses reveal context‐dependent interactions between vertically migrating invasive and native mesopredators and shared prey

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Variability and possible adaptive significance of day-time vertical distribution ofLeptodora kindtii (Focke) (Cladocera) in a shallow eutrophic lake

The day-time vertical distribution of different size classes ofLeptodora kindtii in Tjeukemeer was studied during two contrasting years: 1978 and 1979. During the first year (1978) the predation pressure exerted by the planktivorous fish was rather high, whilst this was ca. 15 times lower during 1979. The diel vertical migration of different size classes ofLeptodora, was studied twice during 1979. Both studies support the predator avoidance hypothesis and also emphasize the idea that diel vertical migration is a dynamic rather than a fixed behaviour trait within a population.     

Diel variation in availability and vulnerability of fish to a survey trawl

The effect diel variation in availability and vulnerability has on the fish capture process of a groundfish survey trawl was examined from data collected during an experiment measuring fish escapement underneath the footgear. The survey trawl was significantly more efficient at night in capturing American plaice, Hippoglossoides platessoides, and yellowtail flounder, Limanda ferruginea, but no difference was found in catches of Atlantic cod, Gadus morhua. This increase in capture efficiency at night is related to an increase in availability and vulnerability associated with diel variation in light intensity. No evidence of diel vertical migration was found and changes in availability were linked to visual dependent gear avoidance by the fish. Regardless of light conditions, the survey trawl had a very low capture efficiency for juveniles of the three species studied. Escapement of this size group was extremely high during both day and night periods due to a fast towing speed, use of large bobbin gear, and possible mesh selection.     

Habitat segregation and interactive effects of multiple predators on a prey assemblage

1. In a field experiment we examined the interactive effects of two common predators of zooplankton, bluegill sunfish (Lepomis macrochirus) and Chaoborus spp. on the growth rate and habitat use of three congeneric prey species (Daphnia). Bluegill and Chaoborus both consume Daphnia, but bluegill also prey on Chaoborus. The prey species, Daphnia pulicaria, D. rosea and D. retrocurva, differed in body size and vertical distribution. We expected the largest species, D. pulicaria, to be most vulnerable to fish predation and the smallest species, D. retrocurva, to be most vulnerable to Chaoborus predation. 2. As we expected, the population growth rate of D. pulicaria was significantly reduced by fish. However, Chaoborus also significantly reduced the growth rate of this species. No significant interaction effect was detected, indicating that the effect of these predators was additive. The growth rates of D. rosea and D. retrocurva were significantly reduced by Chaoborus, but a significant interaction effect indicated that the effect of Chaoborus was stronger in the absence of fish than when fish were present. Therefore the impact of Chaoborus and fish on D. rosea and D. retrocurva was non-additive. The interactive effect of the two predators on D. retrocurva was greater in magnitude than on D. rosea. 3. In the absence of predators, the three Daphnia species showed no differences in mean habitat depth between day and night. Both predators significantly affected diel habitat use of D. pulicaria and D. rosea. Fish caused both of these Daphnia species to move deeper during the day, whereas Chaoborus caused Daphnia to move into shallower water at night. Daphnia retrocurva tended to migrate upwards at night in all predator treatments, but no significant differences in migration were observed among the predator treatments. The effects of predators on habitat use were not interactive for any prey species. 4. Our results suggest that body size, habitat use and the diel migratory response to predators are important factors mediating the interactive effects of multiple predator types on zooplankton.     

Direct and indirect effects of predation by Saduria entomon (Isopoda) on the size-structure of Monoporeia affinis (Amphipoda)

We performed a 6-month laboratory experiment to investigate the direct and indirect effects of predation by the benthic invertebrate predator Saduria entomon on the growth and survival of Monopreia affinis prey individuals in different age-cohorts at low and high prey densities. The experimental results were compared with changes of growth and abundance in corresponding age-cohorts of M. affinis at sites with different S. entomon and M. affinis densities in the deep sublittoral zone of the Bothnian Sea during the same year. In the experiment, the presence of S. entomon reduced growth rate of M. affinis in the 1-year and 2-year age-cohorts at low amphipod density. Increased refuge use by M. affinis, expressed as a decrease in swimming activity in the presence of S. entomon, is suggested to have reduced feeding rate and therefore growth of the amphipods. The recruitment of M. affinis offspring was reduced in presence of S. entomon. In the field, the growth rate of amphipods in the 1-year cohort increased with increasing S. entomon density at low amphipod density. We found no corresponding increase of M. affinis growth in the 2-year cohort. The positive effect on 1-year amphipod growth indicated that predation reduced intra-cohort competition of M. affinis and increased growth of surviving prey at high predator density. In both the experiment and the field data we found indications of size-selective predation on smaller M. affinis specimens. This was because of the changed ratio between number of individuals in the juvenile age-cohorts and lower recruitment of amphipod offspring connected to S. entomon density. The experimental results and field data suggest that predation by S. entomon may have both direct and indirect effects on the size-structure of M. affinis populations. Received: 22 June 1998 / Accepted: 15 March 1999     

Pre‐dispersal seed predation of bayberry Myrica rubra by Thiotricha pancratiastis (Lepidoptera: Gelechiidae) on Yakushima Island,Japan

We investigated pre‐dispersal seed predation by insects in a bayberry Myrica rubra Sieb. et Zucc. (Myricaceae) on Yakushima Island, Japan. To clarify the patterns of seed fate and predation, all fruit that fell into seed traps were collected to allow any insect larvae within the fruit to emerge, and the fruit were finally dissected to determine whether or not they had been attacked by insect predators. Two lepidopteran species, Thiotricha pancratiastis (Meyrick) (Gelechiidae) and Neoblastobasis spiniharpella Kuznetzov & Sinev (Blastobasidae), emerged from the fruits. Thiotricha pancratiastis is the major seed predator of M. rubra, attacking the fruits intensively during the primary stage of fruit development. Thiotricha pancratiastis had been known as a foliage feeder (leaf miner) of M. rubra, but we revealed that the insect is also an important seed predator of the bayberry.     

Modification of the diel vertical migration of Bythotrephes longimanus by the cold-water planktivore, Coregonus artedi

1. The weak diel vertical migration observed in the large cladoceran Bythotrephes longimanus seems contradictory to the predator‐avoidance hypothesis that predicts large zooplankton should have long migration amplitudes. However, cold‐water planktivores, especially Coregonus spp., are a main source of mortality for Bythotrephes and hence a deeper migration would result in a greater overlap with these hypolimnetic planktivores. We hypothesized that Coregonus artedi (cisco) modifies the normal vertical migration pattern of Bythotrephes, such that the latter stays higher in the water column during the day and thus migrates less extremely at night. 2. The vertical distribution of Bythotrephes during the day was determined from single visits to six lakes in Ontario, Canada, all of which contain warm‐water, epilimnetic planktivores but differing in whether they contain cisco. One lake of each fish type was sampled day and night every 2–3 weeks over the ice‐free season to examine daytime depths and migration amplitude. 3. The vertical migration of Bythotrephes differed in the presence and absence of cisco. In the lakes with cisco, there were significantly fewer Bythotrephes in the hypolimnion and they were higher in the water column during the day. Migration amplitude was smaller in the cisco than in the non‐cisco lake. These observations were not attributable to differences in physical factors, and, although not conclusively attributable to cisco, are consistent with an effect of cisco. 4. We suggest that diurnal depth selection by Bythotrephes in lakes containing cisco is a trade‐off between the risk of predation by warm‐ versus cold‐water predators, balanced by the benefits of increased temperature and feeding rates near the surface. Even in lakes without cisco, however, the vertical migration of Bythotrephes was less than expected, suggesting that diurnal depth selection is a balance between the risk from warm‐water planktivores and access to sufficient light to feed effectively.