Formation and maintenance of aggregations in walleye pollock,Theragra chalcogramma,larvae under laboratory conditions: role of visual and chemical stimuli

Synopsis Although planktonic marine fish larvae are often distributed in aggregations, the role of behavioral responses to environmental factors in these aggregations is not well understood. This work examines, under laboratory conditions, the influence of visual and chemical stimuli in the formation and maintenance of aggregations in walleye pollock,Theragra chalcogramma, larvae. Larvae were exposed to a horizontal gradient of light (visual stimulus), prey scent (chemical stimuli: squid/copepod and rotifer) or prey density (visual & chemical stimuli: rotifers). While larvae did not respond to prey scent, they did respond to a gradient of light or prey, which resulted in the formation and maintenance of aggregations. Larvae moved into and remained in a zone of higher light intensity (0.56 versus 0.01 mol photons m^-2 s^-1). Once encountering a patch of prey, larvae remained aggregated within the patch to feed. In nature, movement of walleye pollock larvae in response to selected environmental factors (e.g., gravity, light, temperature, turbulence) may serendipitously bring them into contact with prey patches, where they then could remain to feed as long as light intensity remained at or above levels necessary for feeding.     

Sediment-associated cues for larval settlement of the infaunal spionid polychaetes Polydora cornuta and Streblospio benedicti

Do patchy distribution patterns of infaunal polychaetes result from active site selection of larvae influenced by sediment-associated microbial cues? This hypothesis was tested with still-water laboratory settlement assays revealing the acceptance or rejection of polychaete larvae to qualitatively different sediments. Laboratory brood cultures of the spionid polychaetes Polydora cornuta and Streblospio benedicti yielded a sufficient number of larvae with planktotrophic development for bioassays. High settlement rates (75-95%) of test larvae were observed in response to natural sediment. Sterilization of natural sediment significantly decreased settlement of P. cornuta (25-55%) while combustion of sediment significantly decreased the settlement rate in both species (5-50%). Differences in settlement responses to sediments treated by sterilization or combustion most likely resulted from a variety of factors such as modified sediment fabric, grain size distribution and quantity of adsorbed organic matter. To experimentally address the potential role of microorganisms and microbial metabolites as mediators of larval settlement, ashed sediment was inoculated with viable microorganisms obtained from natural sediment. In both polychaete species, this treatment significantly increased larval settlement in comparison to the control of ashed sediment indicating that larval settlement was at least partially mediated by the presence of microorganisms associated with sediment.     

Microhabitat and rhythmic behavior of tiger beetle Callytron yuasai okinawense larvae in a mangrove forest in Japan

Mangrove forests are regularly flooded by tides at intervals of approximately 12.4 h (tidal rhythm). Larvae of the tiger beetle Callytron yuasai okinawense in a mangrove forest made shallow burrows in mounds up to 1 m in height constructed by the mud lobster Thalassina anomala. No larval burrows were observed on the forest floor, which was very muddy even during low tide. Some larvae plugged the burrow openings before they were submerged at high tide. The mean interval between consecutive burrow plugging events was 12.37 h, which is similar to the period of tidal cycles. Nine out of 30 larvae plugged the burrow openings even when the burrows did not become submerged. Plugging behavior may be governed by an endogenous biological clock, or may be a response to exogenous information about tidal level (e.g. moisture seeping through the ground).     

Strategies of burrowing in soft muddy sediments by diverse polychaetes

Muddy sediments are elastic solids through which morphologically diverse animals extend burrows by fracture. Muddy sediments inhabited by burrowing infauna vary considerably in mechanical properties, however, and at high enough porosities, muds can be fluidized. In this study, we examined burrowing behaviors and mechanisms of burrow extension for three morphologically diverse polychaete species inhabiting soft muddy sediments. Worms burrowed in gelatin, a transparent analog for muddy sediments, and in natural sediments in a novel viewing box enabling visualization of behaviors and sediment responses. Individuals of Scalibregma inflatum and Sternaspis scutata can extend burrows by fracture, but both also extended burrows by plastic deformation and by combinations of fracture and plastic deformation. Mechanical responses of sediments corresponded to different burrowing behaviors in Scalibregma; direct peristalsis was used to extend burrows by fracture or a combination of plastic deformation and fracture, whereas a retrograde expansive peristaltic wave extended burrows by plastic deformation. Burrowing speeds differed between behaviors and sediment mechanical responses, with slower burrowing associated with plastic deformation. Sternaspis exhibited less variability in behavior and burrowing speed but did extend burrows by different mechanisms consistent with observations of Scalibregma. Individuals of Ophelina acuminata did not extend burrows by fracture; rather individuals plastically deformed sediments similarly to individuals of the related Armandia brevis. Our results extend the range of natural sediments in which burrowing by fracture has been observed, but the dependence of burrow extension mechanism on species, burrowing behavior, and burrowing speed highlights the need for better understanding of mechanical responses of sediments to burrowers.     

Effects of burrow condition and seed handling time on hoarding strategies of Edward's long-tailed rat (Leopoldamys edwardsi)

Many hoarding rodents use burrows not only for dwelling and protection from natural enemies, but also for food storage. However, little is known how burrows used by scatter-hoarding animals influence their foraging behaviors. In addition, handling time for a given food item has a fundamental impact on hoarding strategies of these hoarding animals: food items with longer handling time are more likely to be hoarded due to increasing predation risk because the animals spend more time outside their burrows if they consumed such food. By providing with two types of artificial burrows (aboveground vs. underground) and two types of food items (i.e. seeds) with contrasting handling times, we investigated how burrow condition and handling time co-influence hoarding strategies of a key scatter-hoarding rodent, Edward's long-tailed rat (Leopoldamys edwardsi) in large enclosures in southwest China. We found that only a few animals larder-hoarded fewer seeds when only aboveground burrows were available, while over 80% of the animals preferred to use the underground burrows and hoard significantly more seeds in the burrows when both aboveground and underground burrows were provided simultaneously. We also found that seed handling time significantly affected hoarding strategies of the animals: they consumed and/or scatter-hoarded more Camellia oleifera seeds with shorter handling time outside the burrow, but consumed and larder-hoarded more Lithocarpus harlandii seeds with longer handling time in underground burrows. Our study indicates that both burrow types and seed handling time have important impacts on hoarding strategies of scatter-hoarding animals.     

The impact of polychaete (Nereis virens Sars) burrows on nitrification and nitrate reduction in estuarine sediments

The influence of Nereis virens Sars burrows on nitrification, denitrification and total nitrate reduction was assessed in poor (0.7% organic matter) and rich (2.0% organic matter) sediment from the estuary, Norsminde Fjord. The experiments were performed as assays of potential activity, since natural conditions proved impossible to simulate in the unpredictable burrow environment. The measurements were made in two microprofiles, extending 15 mm into the sediment from the surface and from the burrow wall lining. Both sediment types showed higher potential nitrification in the wall linings than in the surface sediment. This was positively correlated with the content of silt + clay particles and organic matter (i.e. the mucous lining of burrow walls). An elevated nitrate reduction activity was evident in the oxic layer of surface sediment. No such activity pattern was observed in the burrow walls. Denitrification accounted for 27–53 % of the total nitrate reduction. An empirical relationship between the ratio of predicted oxygen penetration into surface and wall sediment and the contribution of nereid burrows to bulk actual nitrification and nitrate reduction is presented. The burrow contribution to bulk nitrification was, in contrast to bulk nitrate reduction, very sensitive to variable oxygen penetrations. Thus, possible short-time changes in nitrate exchange across the wall lining will apparently be regulated by changes in nitrification activity rather than nitrate reduction activity.     

Artificial larviposition sites for field collections of the puparia of tsetse flies Glossina pallidipes and G. m. morsitans (Diptera: Glossinidae)

Tsetse flies Glossina pallidipes Austen and G. morsitans morsitans Westwood deposit their larvae in warthog burrows, in August-November, at Rekomitjie Research Station, Zambezi Valley, Zimbabwe. Artificial burrows, made from 200-l steel drums, were used to sample these flies and to collect their puparia. Sand-filled plastic trays in the burrows served as a substrate for larval deposition. The sand was covered with c. 2 cm of leaf litter after it was shown that only 3% of larvae were deposited on bare sand if both substrates were available. Other burrow modifications - artificially shading the burrow entrance, increasing the relative humidity inside the burrow, or reducing the size of the burrow entrance - significantly decreased deposition rates. The use of burrows in the hot season results in a reduction in the temperature experienced by the puparium towards an assumed optimum level of 26 degrees C. Artificial burrows maintained a mean temperature of 28.5 degrees C during October-November 1998, c. 2.5 degrees C cooler than ambient; earlier work has shown that natural burrows can be c. 5 degrees C cooler than ambient at these times. This may explain why natural burrows in full sunlight were used for larviposition, whereas artificial burrows were used only when they were in deep shade, and why significantly higher proportions of G. pallidipes were found in natural (66%) than in artificial burrows (34%). Better-insulated artificial burrows might produce more puparia with higher proportions of G. pallidipes. Burrows become waterlogged during the rains and may be too cool for optimum puparial development during the rest of the year. The percentages of G. m. morsitans in catches of females from artificial burrows, refuges and odour-baited traps were 34, 26 and < 10% respectively. Traps are biased in favour of G. pallidipes; artificial burrows may show a bias in favour of G. m. morsitans that is a function of temperature. Artificial warthog burrows provide a convenient way of studying the puparial stage in tsetse and for the first time facilitate the capture of females as they deposit their larvae.     

Hidden burrow plugs and their function in the tiger beetle, Cosmodela batesi (Coleoptera, Cicindelidae)

Tiger beetle larvae excavate and live in underground burrows, whose openings they sometimes plug with soil. This study documents the burrow plugging behavior of the tiger beetle, Cosmodela batesi (Fleutiaux), in the field. We also tested the function of burrow plugs in the laboratory. In the field, C. batesi more frequently made a plug when it rained. Most larvae made plugs inside their burrows (rather than at the soil surface), and the use of an endoscope was necessary to detect these sub-surface plugs. In the laboratory, flooding was simulated by artificially introducing water into specially-made arenas. Water filled the entire burrow when there was no plug, whereas plugged burrows maintained air chambers inside. When a plug was broken with a wire, burrows filled up with water. The burrowing and plugging behavior described in this study is likely an important adaptation of C. batesi to its habitat.     

Burrow morphology and utilization of the goby (Parapocryptes serperaster) in the Mekong Delta,Vietnam

Some fish species living in mudflats construct burrows for dwelling and hiding. The goby Parapocryptes serperaster is a burrowing fish in mudflats of many estuaries in South East Asia. This study was carried out in the Mekong Delta, Vietnam, to examine burrow morphology and usage by this species. Morphology of the burrows constructed by P. serperaster was investigated by resin castings in situ to obtain the physical structure and configuration of each burrow. Fish from the burrows were caught and measured before burrow casts were made. Fish burrows comprised several openings, a few branching tunnels and multi-bulbous chambers. The surface openings were circular, and the shapes of branching tunnels were nearly round. The burrows had interconnected tunnels and various short cul-de-sac side branches. The burrow structure differed between fish sizes, but burrow dimensions were positively correlated with fish size, indicating that larger fish can make larger and more sophisticated burrow. The burrow structure and dimensions were not different between the dry and wet seasons. Laboratory observations showed that P. serperaster used body movements to dig burrows in the sediment. Burrows could provide a low-tide retreat and protection from predators, but were not used for spawning and feeding for this goby species. This study indicates that the burrowing activity of gobies is an important adaptation for living in shallow and muddy habitats.     

Influence of sediment characteristics on density and distribution of Ocypodoid crab burrows (superfamily: Ocypodoidea) along the coastal areas of Pakistan

Crabs belong to the superfamily Ocypodoidea are a significant component of benthic fauna and considered as ecosystem engineers because of their dynamic role as an active burrower in mangrove and estuarine environment. The current investigation was to evaluate the crab burrow density, diameter and total area of burrow opening along the coast of Pakistan. The variations in burrow properties and their relation to sediment characteristics were also evaluated to recognize the most influencing variables of sediments that effects on crab burrows. All crab burrow and sediment characteristics differed significantly (p < 0.05) among the monitoring sites. Regression analysis showed that crab density was significantly correlated with burrow density (P < 0.001). Moreover, burrow density was noticed significantly greater (p < 0.05) than crab density. Pearson correlation analysis reveals that moisture, porosity, organics, sand and mean grain size observed as most influencing the features of sediment to determine the ecological functioning of crab burrows in mangrove and mudflats of Pakistan.     

The retention of learning during metamorphosis of the crested newt (Triturus cristatus)

Larvae of the urodeles, Triturus cristatus, can be taught a simple discrimination task for food reward. The preference established by training is retained by the adult newts after metamorphosis. No significant change in the natural preference of these animals occurs after metamorphosis. Adult animals appear to be more difficult to train than the larvae.     

The effects of temperature and salinity on ventilation behavior of two species of ghost shrimp (Thalassinidea) from the northern Gulf of Mexico: a laboratory study

Thalassinidean shrimp are among the most important bioturbators in coastal ecosystems. The species Lepidophthalmus louisianensis and Callichirus islagrande are found in dense aggregations (up to 400 burrows m⁻²) along sandy and muddy shores of the northern Gulf of Mexico. These shrimp actively ventilate their burrows to provide oxygen and eliminate wastes. In doing so, they expel nutrient-rich burrow water to the overlying water column, potentially altering nutrient cycling and benthic primary productivity. To develop a mechanistic understanding of the role of burrowing shrimp in nutrient processes, we must first examine how changes in environmental conditions alter the frequency, strength, and duration of ventilation. Field measurements of burrow temperature and salinity suggest that the burrow serves as a buffer from the highly variable conditions found in these estuarine, intertidal habitats. Temperatures at sediment depths >30 cm were generally warmer in winter and cooler in summer than at the sediment surface. Burrow salinities, measured at low tide, were consistently higher than adjacent open water. We used these measurements to parameterize laboratory studies of burrow ventilation in artificial burrows made of plastic tubing and in more natural sediment mesocosms, and studies of oxygen consumption in small glass containers. Rates of oxygen consumption and burrow ventilation by L. louisianensis were lower than those of C. islagrande, perhaps reflecting a lower overall activity rate in the former species which resides in less permeable sediments. Generally, increased temperature had a significant positive effect on oxygen consumption for both species. Salinity had no effect on oxygen consumption by L. louisianensis, reflecting the ability of this species to exist in a wide range of salinities. In contrast, oxygen consumption rates of C. islagrande, which is less tolerant of low salinity, were significantly higher at 35‰ than at 20‰. Ventilation rates were highly variable, and shrimp in artificial burrows tended to have consistently higher ventilation rates than those in sediment mesocosms. There is a trend toward more frequent ventilation at 30 °C for both species. Salinity had no effect on ventilation for either species. Our results suggest that thalassinideans exhibit highly variable and species-specific ventilation patterns that are more likely to be affected by temperature than salinity. Increased ventilation at higher temperatures seems to coincide with increased oxygen consumption at these temperatures, although a similar finding was not made for salinity treatments.     

Distribution,burrowing, and growth rates of the clam Tridacna crocea on interior reef flats

Summary Larvae of the burrowing clam Tridacna crocea (Tridacnidae) settle preferentially on top of detached coral heads lying on the surface of the interior reef flat in the Great Barrier Reef province. This species burrows as it grows, eroding the central top surfaces of coral boulders, producing structures that superficially resemble micro-atolls. Storm surges roll these coral heads onto the now flattened surface, killing the live population of clams, and exposing the fresh underside for unimpeded larval settlement. As these clams grow and burrow into the substratum, the coral head becomes progressively flattened and finally breaks apart. Field observations and growthring data documented growth rate; growth rates plus burrow volumes were converted to annual sediment production. At average population densities approximately 140 gm/m²/yr of coral are eroded. Concomitant with erosion is a calcium carbonate increase in the shell of these clams amounting to 60gm/m²/yr. Assuming a stable population structure, with annual mortality equal to annual estimated growth, total sediment production is 200 gm/m²/yr. Clams are usually aggregated at higher densities, however, with numbers regularly exceeding 100 clams/m². Consequently maximum sediment production rate locally is often 4,500 gm/m²/yr.     

Meandering worms: mechanics of undulatory burrowing in muds

Recent work has shown that muddy sediments are elastic solids through which animals extend burrows by fracture, whereas non-cohesive granular sands fluidize around some burrowers. These different mechanical responses are reflected in the morphologies and behaviours of their respective inhabitants. However, Armandia brevis, a mud-burrowing opheliid polychaete, lacks an expansible anterior consistent with fracturing mud, and instead uses undulatory movements similar to those of sandfish lizards that fluidize desert sands. Here, we show that A. brevis neither fractures nor fluidizes sediments, but instead uses a third mechanism, plastically rearranging sediment grains to create a burrow. The curvature of the undulating body fits meander geometry used to describe rivers, and changes in curvature driven by muscle contraction are similar for swimming and burrowing worms, indicating that the same gait is used in both sediments and water. Large calculated friction forces for undulatory burrowers suggest that sediment mechanics affect undulatory and peristaltic burrowers differently; undulatory burrowing may be more effective for small worms that live in sediments not compacted or cohesive enough to extend burrows by fracture.     

The Use of Conspecific Chemical Cues for Cannibal Avoidance in California Newts (Taricha torosa)

Adult California newts, Taricha torosa (Caudata: Salamandridae), prey on both conspecific egg masses and larvae. We used both laboratory and field experiments to determine if newt larvae use conspecific chemical cues to avoid adult intraspecific predators. Larvae showed significant hiding responses when exposed to adult cues in both the field and the laboratory. Larvae did not exhibit significant hiding responses to chemical cues from invertebrate predators.     

Understanding the toad code: Behavioural responses of cane toad (Chaunus marinus) larvae and metamorphs to chemical cues

Abstract Larvae of many anuran taxa display strong behavioural responses to chemical cues, including alarm signals from injured conspecific tadpoles. We exposed tadpoles and metamorphs from an Australian population of the invasive cane toad (Chaunus[Bufo] marinus) to a range of chemical stimuli and quantified their responses both in the laboratory and in the field. Filtered fluids containing scent cues from crushed conspecifics elicited strong avoidance from tadpoles, whereas other cues (e.g. scent of food, of native‐range fish or urodele predators, and thermal stimuli) did not. Apparent aggregation of tadpoles in response to scent cues proved to be an artifact of tank design, and was an indirect consequence of avoidance of those cues. Field trials confirmed that free‐ranging toad tadpoles and metamorphs avoided chemical cues from crushed conspecifics, suggesting that the chemicals inducing this response might provide an opportunity to develop targeted control methods for this invasive species.     

Behavioral Response of Corophium volutator to Shorebird Predation in the Upper Bay of Fundy,Canada

Predator avoidance is an important component of predator-prey relationships and can affect prey availability for foraging animals. Each summer, the burrow-dwelling amphipod Corophium volutator is heavily preyed upon by Semipalmated Sandpipers (Calidris pusilla) on mudflats in the upper Bay of Fundy, Canada. We conducted three complementary studies to determine if adult C. volutator exhibit predator avoidance behavior in the presence of sandpipers. In a field experiment, we monitored vertical distribution of C. volutator adults in bird exclosures and adjacent control plots before sandpipers arrived and during their stopover. We also made polymer resin casts of C. volutator burrows in the field throughout the summer. Finally, we simulated shorebird pecking in a lab experiment and observed C. volutator behavior in their burrows. C. volutator adults were generally distributed deeper in the sediment later in the summer (after sandpipers arrived). In August, this response was detectably stronger in areas exposed to bird predation than in bird exclosures. During peak predator abundance, many C. volutator adults were beyond the reach of feeding sandpipers (>1.5 cm deep). However, burrow depth did not change significantly throughout the summer. Detailed behavioral observations indicated that C. volutator spent more time at the bottom of their burrow when exposed to a simulated predator compared to controls. This observed redistribution suggests that C. volutator adults move deeper into their burrows as an anti-predator response to the presence of sandpipers. This work has implications for predators that feed on burrow-dwelling invertebrates in soft-sediment ecosystems, as density may not accurately estimate prey availability.     

Circatidal rhythmic behaviour in the coastal tiger beetle Callytron inspecularis in Japan

Larvae of the coastal tiger beetle Callytron inspecularis (W. Horn) (Coleoptera: Cicindelidae) plug their burrow opening before submergence at high tide. Field observations showed that burrow plugging was a rhythmic behaviour that coincided with the tidal cycle (ca. 12.4 h). On average, larvae plugged their burrows 41.8 min before the tide covered the habitat. The mean interval between consecutive burrow-plugging events in the field was 12.40 h. In the laboratory, in the absence of tidal inundation, the mean interval between consecutive burrow-plugging events was 12.45 h. This suggests that the burrow-plugging rhythm of the coastal tiger beetle is governed by an endogenous circatidal rhythm.     

Reactions of <Emphasis Type="Italic">Rana arvalis</Emphasis> Nilss. Tadpoles at Early Developmental Stages to Some Natural Chemical Stimuli

We have investigated behavioral responses of Rana arvalis Nilss. tadpoles to water carrying various water-soluble chemical stimuli: natural lake water; dechlorinated tap water; water in which tadpoles were kept; water with chemical characteristics of the jellylike remains of their own or other clutches after tadpole hatching; boiled nettle, which was used as food for laboratory animals; and water containing toxins of the skin glands of the common toad Bufo bufo L. The preference for natural water over the tap water was revealed, as well as the fact that naive tadpoles were attracted by the “smell” of nettles and tried to avoid toxins.