Distribution,substratum preference and burrowing behaviour of Lovenia elongata (Gray) (Echinoidea: Spatangoida) in the Gulf of Elat ('Aqaba), Red Sea

Lovenia elongata (Gray), the Indo-West Pacific spatangoid, was studied in the Gulf of Elat ('Aqaba) at the northern end of the Red Sea. The occurrence and local distribution of this macrobenthic burrowing species was determined and related to the particle size distribution of the substrata in which it was found. Its habitat preference is for clean, grassless, sandy bottoms in intertidal areas (beaches and sandbars), and on sublittoral level bottoms of protected bays and lagoons sheltered from heavy wave action. The preferred sediments were found to range from medium to fine sand. The population density, size frequency distribution, growth, reproduction, recruitment, and mortality of a sublittoral population of L. elongata (at Wadi Taba, Sinai) were studied.The mode of life of this burrowing heart urchin was observed in situ and in aquaria. Its burrowing and emergence behaviour are described. Experiments were carried out on the effect of particle size on burrowing behaviour. There are significant differences in the ability of L. elongata to manipulate substrata of varying grain size. The burrowing process was found to be most rapid in natural sand of medium-fine composition, corresponding to the normal sediment of the urchin's habitat. Such sediments were also found to be suitable for successful larval settlement; juveniles did not survive in coarser sand.Differences in population density and spatial distribution between various size groups are attributed to differences in their ability to manipulate the substratum. L. elongata is found to be morphologically suited to sand, particularly medium-fine sand; however, it is restricted by its morphology to shallow burrowing, since it lacks specialized mechanisms for deeper burrowing and funnel building.     

A versatile laboratory stream with examples of its use in the investigation of invertebrate behaviour

A versatile and simple laboratory stream was designed and used to investigate the burrowing activity of two insects in response to changes in water velocity and substrate type.Aphelocheirus aestivalis adults were unable to burrow into sand, however, a small proportion of juveniles did burrow in this substrate. The presence of sand in gravel reduced the burrowing success of adults. Steady increases in flow stimulated the burrowing response of both adults and juveniles on gravel and sandy gravel.Ephemera danica was unable to burrow in sand alone at the velocities used in the experiment. The presence of particles greater than 2 mm in diameter in the substrate appeared to be essential for successful burrowing under the test conditions. An increase in flow from 3 to 8 cm s^–1 resulted in an increase in burrowing. The time taken for each specimen to burrow varied widely within replicate tests but most specimens had penetrated the substrate within 150 seconds of introduction. The implications of these observations in influencing the microdistribution of these species are discussed.     

The mechanism of burrowing of Sipunculus nudus

The method of burrowing of Sipunculus nudus L. and the movement beneath the sand is described. Pressures in the coelom have been measured and the forces exerted during burrowing are calculated.     

Effect of Soil Type on Carrot Weevil Movement Behavior

To examine the effect of soil type on carrot weevil, Listronotus oregonensis, (Coleoptera: Curculionidae), movement behavior we conducted a series of laboratory arena experiments with the use of three soil types: mineral soil, organic soil and pure sand. We found that soil type influences carrot weevil burrowing activity and movement and showed that carrot weevil is more inclined to burrow into mineral and organic soils than pure sand and displays avoidance of the latter. While barriers constructed out of pure sand induced burrowing activity only in 3% of tested weevils, mineral and organic soil barriers initiated burrowing in 67% and 38% of tested weevils respectively. All weevils crossed pure sand barriers, whereas 18% (mineral soil) and 6% (organic soil) were arrested and engaged in burrowing activities. In addition, carrot weevil movement behavior at the border of two substrates, revealed high edge permeability of mineral and organic soils and reduced permeability of pure sand edges. Over 70% of weevils released on organic soil ended up not crossing the border with another substrate. All tested weevils released on pure sand crossed the border within less than 1 min. Replication of the experiments with autoclaved soil suggested that the observed differences in carrot weevil movement behavior were largely due to soil texture and not strongly influenced by micro-organisms or odor.     

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.     

Studies on the behaviour and substratum preference in juveniles of the marine prawn,Metapenaeus monoceros (Fabricius)

Circadian rhythmicity in the behaviour of the marine prawn, Metapenaeus monoceros (Fab) and its substratum preference has been observed under laboratory conditions. In the morning at 9. a.m. 80% prawns remain buried and in the afternoon at 3 p.m. 60% burrowing was observed. A contrasting situation was observed in the night i.e. 17% burrowing at 9 p.m. and 30% burrowing at 3 a.m. was registered. In the second set of experiments juveniles were subjected to four types of substrata viz. mud, black fine sand, white coarse sand and stones. It was noted that M. monoceros showed more affinity towards the mud and less preference to other substrata. The obtained results are discussed to provide clues for prawn fishing.This work was supported by I.C.A.R., New Delhi through a project on Reprod. Endocr. Edible prawns.     

Burrowing of the lingulid brachiopod Glottidia pyramidata: its ecologic and paleoecologic significance

Contrary to popular assumption, the pedicle of Glottidia is not its principal burrowing organ. The brachiopod props itself up with the pedicle and enters the sediment with the valves leading, anterior end first. The pedicle trails behind. Burrowing is accomplished by cyclical valve motions: rotary, sliding, and gaping movements are used. Rapid valve closure ejects water from the mantle cavity to loosen the sediment. The lateral setae convey mucus-bound sand posteriorly (upwards). X-radiography shows that the burrows are U-shaped: in a few hours, the animal reappears in feeding position with the anterior end pointing out of the sediment and the pedicle extending down into the burrow. This burrowing process explains the substrate preferences of lingulids. The thick. closely spaced setae are adapted for burrowing. The spacing between individuals is increased, suggesting competition for food.     

Parasitism and the burrowing depth of the beach hopper Talorchestia quoyana (Amphipoda: Talitridae)

Talitrid amphipods spend their days burrowed in sand to avoid predators as well as desiccation and heat stress, although other factors may influence burrowing depth. We investigated the potential role of mermithid nematode parasites in determining burrowing depth in the amphipod Talorchestia quoyana. Mermithids grow as parasites inside amphipods until they reach adulthood, when they must emerge from their host into moist sand to complete their life cycle and reproduce. When allowed to burrow to a depth of their choice in experimental situations, large amphipods burrowed deeper than small ones. In addition, deep-burrowing amphipods were more likely to be infected by mermithid nematodes, and harboured longer worms, on average, than amphipods that burrowed close to the sand surface. This last result is not an artefact of the larger size of deep-burrowing amphipods: the increase in worm length with increasing depth was found after statistical correction for host size. In other words, amphipods that burrowed deeper harboured longer worms than expected based on their body size, whereas those that stayed near the surface of the sand column harboured worms shorter than one would expect based on host size. This implies that the greater burrowing depth of infected amphipods is a consequence, and not a cause, of infection. These results emphasize the importance of parasitism as a determinant of the small-scale spatial distribution of their hosts.     

The relationship of sand moisture to burrowing depth of the sand-beach isopod tylos punctatus Holmes and Gay

The daytime distribution of Tylos punctatus Holmes and Gay was examined by coring on a sand beach in the northern Gulf of California. Moisture and temperature measurements in the cores indicated that the isopods burrow downward to reach sand with at least 1% moisture. Animals experimentally confined to sand with 0.1% moisture showed high mortality rates. We suggest burrowing depth is primarily controlled by a requirement for saturation humidity in the surrounding sand.     

Life in the arena: infaunal gastropods and the late Phanerozoic expansion of marine ecosystems into sand

Marine ecosystems have expanded into the infaunal realm below the surface of soft sediments throughout the Phanerozoic eon. During the Palaeozoic era, this expansion largely involved sedentary animals living in permanent resting places. Active sand‐burrowing animals colonized the infaunal environment later, but when this happened and when specialization for infaunal life evolved remain open questions. Here, phylogenetic evidence, fossil occurrences and previously established criteria for recognizing the sand‐burrowing habit in marine gastropods are used to determine how many gastropod clades became infaunal and when the transitions from surface‐dwelling to infaunal life occurred. At least 20, and as many as 35, clades (all but one of post‐Palaeozoic age) contain actively infaunal species. The overwhelming majority (15 of 20 clades) became infaunal during the Cenozoic, and clades with hundreds of infaunal species in the living fauna diversified beginning in the Early Miocene. The repeated evolution of, and specialization to, the sand‐burrowing habit by gastropods and other animals was enabled by increased habitat availability and higher marine productivity, and was necessitated by intensifying predation. As a result, the infaunal realm was transformed from a marine refuge to an integrated part of the marine biosphere in which high performance in locomotion and defence has become the norm.     

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.     

Water relations of the burrowing sandhill frog, Arenophryne rotunda (Myobatrachidae)

Arenophryne rotunda is a small (2–8 g) terrestrial frog that inhabits the coastal sand dunes of central Western Australia. While sand burrowing is a strategy employed by many frog species inhabiting Australia’s semi-arid and arid zones, A. rotunda is unique among burrowing species because it lives independently of free water and can be found nocturnally active on the dune surface for relatively extended periods. Consequently, we examined the physiological factors that enable this unique frog to maintain water balance. A. rotunda was not found to have any special adaptation to reduce EWL (being equivalent to a free water surface) or rehydrate from water (having the lowest rehydration rate measured for 15 Western Australian frog species), but it was able to maintain water balance in sand of very low moisture (1–2%). Frogs excavated in the field were in dune sand of 4.4% moisture content, as a consequence of recent rain, which was more than adequate for these frogs to maintain water balance as reflected by their low plasma and urine osmotic concentrations. We suggest that in dry periods of the year, A. rotunda can achieve positive water balance by cutaneous water uptake by burrowing deeper into the substrate to where the percent water content is greater than 1.5%.     

Studies of the biology of the lesser weever fish Trachinus vipera Cuvier

The anatomical adaptations to a benthic mode of life are reviewed and discussed. The functioning of the pectoral, pelvic and anal fins in moving sand from underneath the animal, during burrowing, are described. Jets of water directed downwards from the gill chambers, produce a fluid sand-water mixture which facilitates the functioning of the pelvic fins, and the movement of the head into the sand. The direction of water movements associated with gill ventilation, which are propelled almost exclusively by the branchiostegal rays, are outlined. Inhalation is normally through the mouth, which is protected by papillae, from being blocked by sand. When the mouth is covered with sand, the water flow through it is reduced, and some inhalation occurs through the dorsal margin of the operculum, involving use of the supracliethrum.     

Response of Citrus Seedlings to Radopholus similis in Two Soils

Radopholus similis was less pathogenic to greenhouse-grown citrus seedlings in Leon loamy sand than in Lakeland fine sand. This was not affected by different watering regimes. Seedling growth reduction by the burrowing nematode in either soil, compared with noninfected controls, was significant at the 1% level.     

Burrowing abilities and swash behavior of three crabs, Emerita analoga Stimpson, Blepharipoda occidentalis Randall, and Lepidopa californica Efford (Anomura, Hippoidea), of exposed sandy beaches

To investigate factors related to the distribution of intertidal species, and specific predictions of the swash exclusion hypothesis for exposed sandy beaches, we compared the burrowing abilities and swash behavior of three species of anomuran crabs in the superfamily Hippoidea (Emerita analoga, Blepharipoda occidentalis and Lepidopa californica) which commonly inhabit the intertidal and shallow subtidal zones of beaches along the California coast. Burrowing times in the laboratory increased significantly with crab size for all species in five sediment grain sizes ranging from fine sand to gravel (0.15 to 3.24 mm). For each species, burrowing times differed significantly among sand grain sizes, ranging from 0.3 to 21.5 s. Burrowing times for the hippid crab, E. analoga, were relatively constant across sediment types, while those of the albuneid crabs, B. occidentalis and L. californica, were rapid in fine to medium sands, and much slower in coarser sediments. Our results indicate that E. analoga is a substrate generalist while L. californica and B. occidentalis are substrate sensitive. Pre-burrowing times and behavior, distance moved, and burrowing times differed among the species in the swash zone. Combined times of preburrowing and burrowing were shorter than the swash period (6 s) for most E. analoga individuals. Fifty percent of the individuals of L. californica reached the substrate and burrowed in the swash period, while no individuals of B. occidentalis burrowed in that time. Pre-burrowing behavior and time may be valuable in explaining spatial and temporal patterns in the distribution of hippoid crabs on California beaches. Our results support predictions of the swash exclusion hypothesis concerning the burrowing and locomotory abilities of sandy beach macrofauna. The substrate generalist characteristics, and unique orientation and swimming abilities of the hippid crab, E. analoga, in intertidal swash may help explain the success of this species and its congeners, and have important implications for understanding patterns of macrofauna community structure on exposed sandy beaches in California and other regions.     

Burrowing of Priapulus caudatus (Vermes) and the significance of the direct peristaltic wave

Priapulus caudatus is an active predatory animal found burrowing in soft mud. It may employ relatively high body fluid pressures during escape reactions (up to 6 kPa recorded) but normal burrowing probably involves much lower pressures (< 2 kPa). Three points d'appui are formed during the burrowing cycle. A characteristic direct peristaltic wave (DPW) occurs at a low pressure stage in each cycle (<0–5 kPa). The DPW is regarded as an adaptation to burrowing in soft mud. Its function appears to be to transfer body wall anteriorily, providing "slack", which is used to allow the trunk to extend forward into the cavity vacated by retraction of the large praesoma at the end of each cycle. The circular and longitudinal muscles of the posterior trunk act synergically through most of the burrowing cycle.     

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.     

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.     

Water pumping and analysis of flow in burrowing zoobenthos: an overview

Burrowing animals maintain contact with the water above the sediment by pumping water through a tube system and therefore measurements of water pumping rate of burrowing animals is of crucial importance for the study of many processes both within and above the sea floor. This review deals with the measuring of water pumping and the analysis of flow generated by burrowing deposit- and filter-feeding zoobenthos in order to determine the type of pump and mechanisms involved, flow rate, pump pressure, and pumping power. The practical use of fluid mechanical principles is examined, and it is stressed that not only the pump pressure that a burrowing animal can apply is of interest for assessing the energy cost of pumping, but also the distribution of excess pressure along its burrow is of importance for assessing the seepage flow of oxygen-rich water into the sediment surrounding the burrow because this bioirrigation exerts a considerable effect on the chemistry and microbiology of sediments. Dense populations of burrowing filter-feeding zoobenthos also interact with the water above the sediment interface and this is reflected in the development of phytoplankton concentration profiles above the filter-feeding animals. In stagnant situations the near-bottom water may be depleted of food particles, depending on the population filtration rate and the intensity of the biomixing induced by the filtering activity. But moderate currents and the biomixing can presumably generate enough turbulence to facilitate mixing of water layers at the sea bed with the layers above where food particle concentrations are relatively higher. Following a brief summary of types of burrowing benthic animals, common methods for measuring pumping rates are described along with examples. For estimating the required pump pressure, biofluid mechanical theory for flow in tube–pump systems is summarised (elaborated in Appendix A). Specific examples are given to illustrate general principles and to give an idea of typical values of flow rate, pressure drop and power involved. Finally, some flow effects generated by burrowing animals in and above the sediment are described.     

Infaunal hydraulics generate porewater pressure signals

Many activities by infauna, including burrowing and feeding, involve hydraulic mechanisms. We expected these activities to generate low-frequency pressure waves that would propagate through sediments and be detectable at some distance from the source. Pressure sensors in intertidal sediments recorded large-amplitude porewater pressure signals. Laboratory recordings of single individuals allowed us to identify characteristic signals of arenicolid and nereidid polychaetes and tellinid bivalves. In the bivalve Macoma nasuta, these high-amplitude signals were associated with burrowing, expulsion of pseudofeces, and siphon relocation. In the polychaetes Neanthes brandti and Abarenicola pacifica, the high-amplitude pressure signals were associated with burrowing, burrow construction, burrow ventilation, and defecation. These signals were detectable in the field at distances of at least 20 cm. Since the waveforms are species-specific as well as activity-specific, they may provide a mechanism for prey detection, for predator avoidance, for competitor detection, and perhaps even for mate detection.