Burrowing behaviour of the softshell clam (Mya arenaria) following erosion and transport

Erosion and transport of juvenile individuals may alter the distribution pattern of intertidal bivalves. The burrowing success of recently transported juvenile softshell clams (Mya arenaria) was studied in a laboratory flume under a wide range of hydrosedimentary environments. Juvenile individuals (5-20 mm) were observed under a simulated 30 min slack tide before initiating the flow for a period of 60 min. Five different free-stream velocities (0, 3, 5, 10 and 24 cm s^− 1) and four sediment types (mud, sandy-mud, sand and gravel) were used. The mean proportion of juvenile clams that initiated (MPI) or completed (MPC) a burial decreased with increasing shell length. Erosion from the sediment was more important in large juveniles suggesting that large juveniles may have more difficulty successfully relocating once transported. The MPI increased with increasing flow speed in experimental runs held at speed < 24 cm s^− 1. This was observed in all sediment types. Most individuals were unable to burrow at 24 cm s^− 1 because they got eroded. The MPC also increased with increasing flow speed in mud, sandy-mud and sand. The MPC's response to flow was more complex in gravel because of a shell length × flow speed interaction effect. Our observations suggest that water movement may induce the burrowing behaviour of recently eroded juvenile clams. Results are discussed in an ecological and aquacultural context.     

Burrowing behaviour of the European eel (Anguilla anguilla): Effects of life stage

The European eel (Anguilla anguilla) is a fascinating species, exhibiting a complex life cycle. The species is, however, listed as critically endangered on the IUCN Red List due to an amalgam of factors, including habitat loss. This study investigated the burrowing behaviour and substrate preference of glass, elver and yellow stages of A. anguilla. Preference was determined by introducing eels in aquaria with different substrates and evaluating the chosen substrate for burrowing. In addition, burrowing was recorded using a camera in all substrate types and analysed for kinematics. The experiments showed that all of these life stages sought refuge in the sediments with particle sizes ranging from sand to coarse gravel. Starting from a resting position, they shook their head horizontally in combination with rapid body undulations until half of their body was within the substrate. High-speed X-ray videography revealed that once partly in the sediment, eels used only horizontal head sweeps to penetrate further, without the use of their tail. Of the substrates tested, burrowing performance was highest in fine gravel (diameter 1–2 mm; lower burrowing duration, less body movements and/or lower frequency of movements), and all eels readily selected this substrate for burrowing. However, glass eels and elvers were able to use coarse gravel (diameter >8 mm) because their smaller size allowed manoeuvring through the spaces between the grains. Further, burrowing performance increased with body size: glass eels required more body undulations compared to yellow eels. Interestingly, the urge to hide within the sediment was highest for glass eels and elvers. Documentation of substrate preference and burrowing behaviour of A. anguilla provides new information about their potential habitat use. Considering that habitat alterations and deteriorations are partly responsible for the decline of the eel, this information can contribute to the development of more effective conservation measures.     

Behavior of the burrowing shrimp Alpheus macellarius in varying gravel substrate conditions

The behavior of the male snapping shrimp, Alpheus macellarius (Decapoda: Alpheidae), was studied in tank experiments with four levels of gravel content for five consecutive days. Markov-chain type analyses identified significant behavioral states that were grouped into six behavioral classes (Wandering, Surveying, Grooming and Foraging, Resting, Hidden). Significant trends in the durations and frequencies of the classes were found across days and periods, with shrimps mainly wandering and burrowing during the first day. Grooming, foraging and resting became more frequent during succeeding days. Shrimps built burrows in the morning and then groomed and foraged mostly in the afternoon. This periodicity implies a greater priority for the shrimp to construct and maintain a burrow rather than forage. Gravel content did not significantly influence burrowing behavior, but marked variations were noted in burrowing success, burrow structure and stability. Animals in 15% and 25% gravel substrates produced more burrows of greater complexity, which lasted longer than those of the shrimps in 0% and 5% gravel setups. Feeding was mainly sediment scavenging suggesting the predominance of a deposit-feeding trophic mode. Modifications in burrow construction and adaptations in burrowing and feeding indicate the ability of A. macellarius to respond to different environmental conditions.     

Diel activity patterns in Metapenaeus and Penaeus juveniles

Small (5–10.9 mm carapace length), medium (11–15.9 mm), and large (16–20.9 mm) juveniles of Metapenaeus anchistus, Metapenaeus sp., Penaeus monodon and P. merguiensis were stocked individually in glass tanks provided with sand substrate, sea water, artificial bamboo shelter, aeration and food. The seven activity types (recorded for each shrimp hourly for 24 h) were classified as below (burrowing) or above substrate (swimming, walking, stationary, in shelter, feeding and cleaning). Shrimp juveniles exhibited a strong diel periodicity — emergence and activity at night and burrowing in the day. The chi-square test showed that type of activity (above/below substrate) was associated with period (light/dark). Diurnal burrowing was greater among Metapenaeus than Penaeus; inversely, above substrate activities were more frequent for Penaeus species compared to Metapenaeus. Feeding was the major above substrate and nocturnal activity for M. anchistus, Metapenaeus sp. and P. monodon. Only P. Monodon used the shelter consistently. Frequency of the 7 activity types was dependent on juvenile size for Penaeus, e.g., the preference for shelters shifted to burrowing with increase in size in P. monodon. Results are discussed in relation to the importance of mangrove habitats in providing shelter to penaeids, in particular the mangrove-associated P. monodon and P. merguiensis.     

Grain-size controls on the occurrence of bioturbation

Grain size and grain-size related stresses impart a significant influence on the ichnological character of marginal-marine deposits. This is evident on the New Brunswick coastline of the Bay of Fundy, Canada, where three coarse-grained marginal-marine deposits are studied to assess grain-size controls on the occurrence and type of bioturbation. Firm mud and sand substrates exhibit the greatest diversity and density of bioturbation (i.e., bioturbation intensity). The types of organisms colonizing sands and firm-mud substrates are variable; however, the resultant trace assemblages are similar. Thixotropic muds exhibit significantly reduced trace diversity and density relative to firm mud, reflecting the additional stress placed on the organisms by the relatively soupy consistency of the sediment. A significant change in the trace assemblage occurs when sediment caliber passes the gradational sand—fine gravel boundary.Four main conclusions can be drawn from this study. First, for mixed sand and gravel, fine gravel, and coarse-gravel deposits, the degree of bioturbation (diversity ^? density) decreases more rapidly onshore (across the intertidal zone) than is noted in sand or mud deposits. Second, there is a decrease in the degree of bioturbation with increasing grain size for substrates composed of sand-sized and larger clasts. Third, burrows in gravels tend to be lined and/or robust, likely to maintain a stable environment within the burrow. Fourth, in coarse-gravel substrates or substrates with a significant component of coarse gravel, burrows are developed between the clasts and tend to be more permanent structures (than those developed in sand or mud), which are generally continuously occupied.The degree of burrowing noted in these modern gravel deposits contrasts with the relative paucity of biogenic structures reported in conglomerates preserved in the rock record. Based on the intensity of burrowing observed in the gravels, we hypothesize that ancient marginal-marine conglomerates are likely bioturbated, but that these burrows are likely distorted during burial and compaction.     

Spatial configurations generated by motile benthic polychaetes

Micro-spatial patterns of five infaunal polychaete species were investigated on tidal flats in the Wadden Sea (island of Sylt, North Sea). Sediment samples were taken within plots of 4 m² at regular distances or with a multiple-cell corer composed of 144 contiguous units. Most of the configurations observed can be related to the mode of feeding. Individuals of the predatory polychaeteEteone longa do not generate discernible spatial structures.Anaitides mucosa, a carnivoruous scavanger, occurs in marked patches. Juveniles ofScoloplos armiger, a burrowing deposit feeder, live in small aggregates which in turn conglomerate to larger ones. Adults are scattered within larger clusters. High density areas of juveniles and adults do overlap. Another burrowing deposit feeder,Capitella capitata, is even more aggregated. Local attractors may cause clusters of exceptional intensity. Territoriality is exhibited by the tube-dwelling surface deposit feedingNereis diversicolor. The juveniles are restricted to interspaces left by the adults.I acknowledge a grant from the Deutsche Forschungsgemeinschaft (DFG).     

Observations on the clypeasteroid Echinocyamus pusillus (O. F. Müller)

The tiny echinoid Echinocyamus pusillus (O. F. Müller) is equipped with specialized external structures that suit it for a wide variety of environments. Special features include the ability to burrow in sediments of fine sand to shell gravel and to climb vertically.Specimens dredged off the west coast of Scotland were observed in aquaria and with the SEM. E. pusillus is characterized by three kinds of spines, and by two of pedicellariae. In contrast to sand dollars, the spines play a passive rôle in the feeding and burrowing operation, probably retaining a defensive nature as in the regular urchins. It is the podia that are chiefly involved in climbing, burrowing, righting, and probably feeding. Surface ciliary currents transport particles, but not to the mouth; they may have a respiratory or cleansing function. Experimental animals did not burrow in either very fine or very coarse sand, probably because a certain relationship exists between particle weight and podia size.E. pusillus shares behavioural and structural characteristics with regular and irregular urchins. It is not a true sand dollar, but may illustrate an evolutionary stage towards such a form.     

Burrow structure and foraging costs in the fossorial rodent,Thomomys bottae

Summary A model for calculating the energy cost of burrowing by fossorial rodents is presented and used to examine the energetics of foraging by burrowing. The pocket gopher Thomomys bottae (Rodentia: Geomyidae) digs burrows for access to food. Feeding tunnels of Thomomys are broken into segments by laterals to the surface that are used to dispose of excavated soil. Energy cost of burrowing depends on both soil type and on burrow structure, defined by the length of burrow segments, angle of ascent of laterals, depth of feeding tunnels, and burrow diameter. In a desert scrub habitat, Thomomys adjust burrow segment length to minimize cost of burrowing. Observed segment lengths (mean=1.33 m) closely approximate the minimum-cost segment length of 1.22 m. Minimizing energy expended per meter of tunnel constructed maximizes efficiency of foraging by burrowing in the desert scrub. Burrow diameter and cost of burrowing increase with body size, while benefits do not, so foraging by burrowing becomes less enconomical as body size increases. Maximum possible body size of fossorial mammals depends on habitat productivity and energy cost of burrowing in local soils.     

The role of substrate feeding earthworms (Lumbricidae) for bioturbation in a beechwood soil

Summary Burrow formation (burrow length and faeces production) of the substrate feeding earthworm Aporrectodea caliginosa (Savigny) in relation to three temperatures (5, 10 and 15°C) and soil moisture contents (48, 60 and 73% water of dry wt) was determined. Soil moisture content affected the burrowing activity of A. caliginosa only below a distinct threshold (60% water of dry wt). At sufficient moisture an, increase in temperature by 5° C approximately doubled the amount of egesta produced. Another substrate feeding earthworm (Octolasion lacteum (Örley)) showed a similar pattern of burrowing activity.The data for the dependence of the faeces production of A. caliginosa on soil temperature and moisture content were combined with field data. The amount of faeces produced by the population of this earthworm species in a beechwood on limestone was calculated to be about 4.23 kg ha^-1 a^-1. The amount of egesta produced by all substrate feeding species is assumed to exceed 6 kg ha^-1 a^-1, which is equivalent to a soil layer of approximately 9 mm. This group of earthworms is therefore considered to be of greater importance for bioturbation and the formation of mull than previously assumed.     

Palaeocommunity Dynamics and Behavioral Analysis of Conichnus: Bhuj Formation (Lower Cretaceous),Kachchh-India

A palaeocommunity of large Conichnus conicus, a conical, cone-in-cone shaped burrow, created by sea anemones, occurs in medium-grained, crossbedded, well-sorted sandstone in the middle part of the Cretaceous Guneri Member of the Bhuj Formation in India. The trace fossil Conichnus is considered to be a common element of the Skolithos ichnofacies and is interpreted to reflect equilibrium movement in response to substrate aggradation. In the present study, three different varieties of Conichnus conicus are distinguished based on morphology and internal fabric. Community dynamics and burrowing behavior are revealed by inter-burrow relationships, burrow initiation levels and sedimentology. Three types of behavior are envisaged: retrusive equilibrium, protrusive equilibrium response, and escape behavior. Palaeocommunity dynamics show that the tracemakers consisted of only adult organisms that initiated burrows during neap tides and are adapted to feed effectively during weak flow conditions. The occurrence of Conichnus palaeocommunity in the Guneri Member indicates the tidal conditions in a fully marine setting. Results presented herein may aid in the understanding of palaeocommunity dynamics in other shallow marine sequences.     

Nutrition and burrowing energetics of the Cape mole-rat Georychus capensis

Summary At 22°C the resting oxygen consumption of G. capensis is 1.13±0.05 cm³O₂·g^-1·h^-1 (mean± S.E.). In loose sandy soil the burrowing metabolic rate was approximately three times that of resting (3.41±0.19 cm³O₂·g^-1· h^-1). Rate of oxygen consumption while burrowing bears a linear relationship with rate of burrowing. The equation of the regression line describing this relationship was used to construct a model for calculating energy expenditure of burrowing in free-living mole-rats. The diet of G. capensis consists of some green plant material and geophyte corms. The latter has a mean gross energy content of 16.36 kJ·g^-1 dry weight. The digestibility coefficient for captive G. capensis fed on sweet potato, was 97.42±0.41%. Data collected from an excavated burrow system revealed that the total energetic cost of constructing the burrow amounted to 79% of the estimated digestible energy available from geophyte corms in the area. A food store in the same burrow system was sufficient to meet the maintenance requirements of an adult G. capensis, resting at 22°C, for approximately 80–85 days. Soil samples taken at random adjacent to the burrow contained corms with a mean estimated digestible energy value of 2084 kJ per m³ of soil. A comparison of energetic cost of burrowing and randomly available digestible energy in the field suggests that foraging patterns are not random.     

Invertebrate colonization of woody debris in coastal plain streams

We studied colonization dynamics and habitat preferences of macroinvertebrates associated with submerged woody debris in Louisiana coastal plain streams. In an in situ experiment, Branches of magnolia Magnolia grandiflora and water oak Quercus nigra were anchored over sand and gravel substrate at two sites in the Bogue Falaya River, St. Tammany Parish, Louisiana, U.S.A. Branches were removed from the units weekly for 4 weeks and biweekly for 1 month to assess colonization. Invertebrate numbers were highest at weeks three and six on magnolia over gravel habitat, although numbers of the most dominant taxa (Baetidae, Hydropsyche sp., Cernotina sp., Oecetis sp. and Hydroptilia sp.) peaked at different times during the study. Abundances of Hydropsyche sp. and Cernotina sp. were greatest on magnolia branches secured over gravel substrate, whereas Oecetis sp. abundances were greatest on oak branches secured over sand substrate.We also surveyed six streams in southeast Louisiana to examine the relative importance of wood, gravel and sand as substrate for lotic macroinvertebrates. In fall 1997 and spring 1998, we took water samples and measured in situ water quality characteristics at one site on each stream, and then collected submerged wood (surface area about 3051 cm²) and sand and gravel substrate (Surber samples totaling 2800 cm² for each substrate) to determine the density and taxonomic composition of resident macroinvertebrates. Highest numbers of invertebrates were collected in fall and spring from gravel substrate (P=0.0001). Macronychus sp. were more abundant on wood than gravel or sand (P<0.0003) and Hydropsyche sp. was more abundant on fall wood samples (P=0.0001). Analyses of water quality parameters revealed associations between Hydropsyche sp. abundance and potassium (P=0.0070) and specific conductance (P=0.0080), although both parameters exhibited interactions with substrate type (P<0.0144).     

Survival and behaviour of juvenile unionid mussels exposed to thermal stress and dewatering in the presence of a sediment temperature gradient

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Gain of palps within a lineage of ancestrally burrowing annelids (Scalibregmatidae)

Scalibregmatidae is a small annelid family of subsurface deposit feeders in sand or mud, which are generally well adapted to infaunal burrowing. The overall morphology of Scalibregmatidae is very similar, with thick bodies, small parapodia, and no prostomial appendages or short horns. The only exception is members of the genera Axiokebuita and Speleobregma that most frequently inhabit crevices or gravel and possess extensive ventral ciliated palps and globular adhesive pygidium. Character reconstruction using maximum likelihood and Bayesian methods show that ciliated palps and adhesive pygidium are synapomorphies of the Axiokebuita–Speleobregma clade. The most likely transformation series is from horns to ciliated palps, the origin of which correlates with the occurrence of Axiokebuita and Speleobregma in crevices or gravel. The wide spaces among rocks or granules yield high permeability and inertial water flow, preventing deposition of organic matter. Under these flow conditions that differ significantly from those of sand or mud bottom, ciliated palps aid to the collection of suspended particles and an adhesive pygidium provides attachment. With palps being a highly debated character in annelid evolution, it is remarkable that prominent ciliated palps are gained within a lineage of ancestrally nonpalpate annelids, most likely increasing their fitness when colonizing a new environment.     

Responses of <Emphasis Type="Italic">Venerupis philippinarum</Emphasis> (Mollusca: Bivalvia) to Predators and Changes in Hydrodynamics

The rate of burrowing into sand by the bivalve mollusk Venerupis philippinarum was studied under laboratory conditions. It is found that an increase in the intensity of water flow to an average of 1000 ml/min stimulates the burrowing of the mollusk, while higher values suppress burrowing; the threshold values are somewhat different for the young and adult specimens. It is shown that the presence of the starfish Asterias amurensis in the experimental environment decreases the rate of burrowing of V. philippinarum by 10–20% depending upon the sizes of the predator and the potential victim, but the depth of penetration of the mollusks into the sediment increases. It is supposed that in V. philippinarum and other species of bivalve mollusks related to the same life form and characterized by a moderate capability to burrow into the ground, the adaptive strategy of threat avoiding (great wave activity, an approach of a predator) is expressed as burrowing into a depth uncommon for usual situations and waiting. This behavior, to a significant degree, provides the capability of some species to inhabit shallow water areas with poor hydrodynamics inhabited by numerous predators.     

Influence of age and environmental factors on burrow-making behavior of the short-tailed cricket,Anurogryllus muticus (De Geer) (Orthoptera: Gryllidae)

For the short-tailed cricket, Anurogryllus muticus, burrow-making behavior is essential. All nymphal instars construct burrows, but in the adult stage the rate of burrowing behavior is age dependent. Increases in photophase and light intensity stimulate burrowing, and the explicit negative phototaxis is correlated with the cricket's inability to exist under dry conditions. Ingestion of substrate during burrow construction may serve to acquire additional moisture. There is no evidence of burrow recognition, and crickets can construct a burrow when needed. The natural distribution of burrows at the plot investigated on Moorea supports the notion thatA. muticus builds burrows where the preferred food plantAlysicarpus vaginalis is most abundant. By minimizing the traveling distance to food sources when foraging they can retreat to their burrow again.     

Intraspecific competition in Tridacna crocea,a burrowing bivalve

Summary Intraspecific competition for space and light occurred when Tridacna crocea burrowed into coralline substratum of boulders on leeward coral reefs in the central Great Barrier Reef near Townsville, Australia. Intensity of competition was linearly related to clam density. Above about 200 clams/m², all clams physically contacted one another and all shells sustained damage. Mortality in isolated populations due to intraspecific competition was estimated at 40%. Principles of intraspecific competition in plants were tested for applicability to T. crocea populations. Juvenile mortality due to competitive stress was density dependent. Aggregated distributions of one year old clams changed to random or regular distribution of adults. Normal size-frequency distribution for juveniles became skewed for older groups. A bimodal size-frequency distribution of the population was related to selective mortality in 1–3 year old clams. Adult mortality due to crowding was less severe but significant. Growth rates were inhibited by competition. Deformations in morphology resulted from crowding. Intraspecific competition for space and light by adults inhibited recruitment of young. Animal adaptations to reduce mortality under crowded conditions were also important. Larvae aggregated on settling and oriented with posterior ends pointed away from nearest neighbors. Positional alignment within the substratum was selectively advantageous. Burrowing posteriorly was preferential, but anterior and sideways burrowing as well as twisting within the burrow were also observed. Movement within substratum served to reduce local damage to the shell. Proteinaceous deposits secreted through perforations in the shell reduced subsequent damage. T. crocea populations exhibited many animal adaptations that reduced mortality during the first years of life, but as cohorts matured, plant-like patterns of competitive interaction became more significant.     

Bioturbation of Burrowing Crabs Promotes Sediment Turnover and Carbon and Nitrogen Movements in an Estuarine Salt Marsh

Ecological functions of bioturbation in ecosystems have received increasing attention over the recent decades, and crab burrowing has been considered as one of the major bioturbations affecting the physical and chemical processes in salt marshes. This study assessed the integrated effects of crab excavating and burrow mimic trapping on sediment turnover and vertical C and N distributions in a Chinese salt marsh in the Yangtze River estuary. Crab burrowing increased soil water content and the turnover of carbon and nitrogen and decreased bulk soil density. Vertical movement of materials, nutrient cycling and reuse driven by crab burrowing might be obstructed by vegetation (Phragmites australis and Spartina alterniflora communities). The amount of soil excavated by crab burrowing was higher than that deposited into burrow mimics. In Phragmites marshes, Spartina marshes and unvegetated mudflats, net transport of soil to the marsh surface was 171.73, 109.54, and 374.95 g m⁻² d⁻¹, respectively; and the corresponding estimated soil turnover time was 2.89, 4.07 and 1.83 years, respectively. Crab burrowing in salt marshes can mix surface and deeper soil over a period of years, accelerating litter decomposition and promoting the efficient reuse of nutrients by plants. Therefore, bioturbation affects soil physical processes and functioning of ecosystems, and needs to be addressed in ecosystem management.     

Push and bite: trade‐offs between burrowing and biting in a burrowing skink (Acontias percivali)

Trade‐offs are thought to be important in constraining evolutionary divergence, as they may limit phenotypic diversification. Limbless animals that burrow head‐first have been suggested to be evolutionarily constrained in the development of a large head size and sexual head shape dimorphism because of potential trade‐offs associated with burrowing. Here we use an acontiine skink (Acontias percivali) to test for the existence of trade‐offs between traits thought to be important in burrowing (speed and force). As head size dimorphism has been shown to be limited in acontiine lizards, thus suggesting constraints on head size and shape, we additionally explore the potential for trade‐offs between burrowing and biting. Our data show that A. percivali uses a burrowing style different from those previously described for caecilians and amphisbaenians, which relies on the use of extensive lateral and dorsoventral head movements. Our data also show that animals use their entire bodies to generate force, as peak force was determined by total length only. Additionally, both bite force and the time needed to burrow into the substrate were principally determined by relative head width, suggesting a trade‐off between biting and burrow speed. Performance data were indeed suggestive of a correlation between bite force and the time needed to burrow, but additional data are needed to confirm this pattern. In summary, our data suggests that trade‐offs may exist, and may have been of crucial importance in shaping the evolution of head shape in A. percivali, and burrowing lizards more generally. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 91–99.     

Parasite infection and sand coarseness increase sand crab (Emerita analoga) burrowing time

Parasites with indirect life cycles require trophic transmission from intermediate hosts to definitive (vertebrate) hosts. Transmission may be facilitated if parasite infection alters the behavior of intermediate hosts such that they are more vulnerable to predation. Vulnerability to predation may also be influenced by abiotic factors; however, rarely are the effects of parasites and abiotic factors examined simultaneously. The swash zone of sandy beaches is a particularly harsh environment. Sand crabs (Emerita analoga) burrow rapidly in the swash zone to avoid predators and dislodgment. We examined prevalence and abundance of the acanthocephalan parasite Profilicollis altmani in sand crabs, and investigated the synergistic effects of sand grain size (an important abiotic factor), parasite infection, body size and reproductive condition on burrowing speed in females, from three California sites. More heavily parasitized crabs burrowed more slowly, making them potentially more vulnerable to predation by marine bird definitive hosts. Ovigerous females harbored more parasites than non-ovigerous females, but burrowed more quickly. All crabs burrowed slowest in the coarsest sand, and burrowing times increased with repeated testing, suggesting that it is energetically costly. Abiotic and biotic factors influence burrowing, and behavioral variation across sites may reflect the response to natural variation in these factors.