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.     

Neoichnological activities of endobenthic invertebrates in downdrift coastal Ganges delta complex,India: Their significance in trace fossil interpretations and paleoshoreline reconstructions

Neoichnological studies of the downdrift coastal Ganges deltaic region (Ganges Delta Complex) indicate the nature and environmental zonation of the lebensspuren of common endobenthic invertebrates dominated by the brachyuran amphibious crab Family Ocypodidae (Ocypode spp., Ilyaplax pusillus and Uca marionis) and are utilized to interpret analogous trace fossils and paleoshoreline environments. The polycha‐ete Diopatra cuprea, the gastropods Turritella spp., Tele‐scopium telescopium and Cerithidea obtusum, some bivalves and the boring crab Charybdis rostrata also produce diagnostic lebensspuren.

The measured ichnoprofiles reveal the development of coast parallel Uca ‐Turritella (backswamps and salt‐marshes; 1 type burrows), Ocypode ‐ Ilyoplax (backshore to foreshore), Charybdis rostrata (foreshore relict woodground; boring structures), polychaete (middle‐lower foreshore; current ‐ oriented agglutinated burrows) and bivalve ‐ gastropod (lower foreshore; trails) ichnozones. Ocypode ‐ Ilyoplax, irrespective of ontogenic stages and sex, produce I, J, U, Y, and multibranched Y ‐ shaped burrows (juvenile ‐ old in backshore versus young ‐ adult in foreshore) in an orderly fashion. Their burrow density and diversity attain a maximum in backshore and upper foreshore respectively. Exceptionally, high burrow populations produce network burrow systems in the backshore. Juvenile pelletal designs (upper foreshore) and general landward burrow inclination are conspicuous.

The described lebensspuren, having a wide range of ancient analogues, provide supportive evidence in the identification of lithified crab burrows, paleoshoreline environments and paleosealevel fluctuations.     

Ecological Predictors of Range Areas and Use of Burrow Systems in the Diurnal Rodent, Octodon degus

Variation in animal space use patterns may be linked to numerous ecological factors affecting survival and reproduction. We examined the relationship between ecology and above‐ and below‐ground components of space use by Octodon degus, a semi‐fossorial rodent in Chile. We monitored the daytime minimum convex polygon and adaptive kernel range areas of 26 individuals and determined the number of burrow systems used by degus during night‐time radiotelemetry and trapping of burrow systems on two study grids at Rinconada de Maipú, a semi‐arid Matorral in central Chile. We quantified food biomass, soil hardness, distance to overhead vegetative cover, and density of burrow openings at putative nest burrows. Degus living on the grid with more shrub cover had larger range areas than degus living on the grid with less cover. The range areas of degus decreased with increasing distance from overhead vegetative cover. There was a weak (but statistically significant) negative relationship between the number of burrow systems used by degus and the distance to vegetative cover and density of burrow openings at burrow systems. Male and female degus had similar range areas. Our results suggest that overhead cover decreases the risk of predation to male and female degus. Degus probably balance the benefits of numerous burrow openings (reduced predation risk) with time and energy requirements of burrow construction and maintenance. Models of space use that consider the effect of multiple ecological variables should measure different dimensions of space use.     

Burrow Use and the Influence of Ectoparasites in Brants' Whistling Rat Parotomys brantsii

In Brants' whistling rat, Parotomys brantsii, single animals inhabit unusually complex burrow systems containing several nest chambers. We investigated burrow use and examined the effect of ectoparasites on choice of alternative nest chambers by radiotelemetrically monitoring the nocturnal sleeping locations of individual whistling rats, before and after treating them to remove ectoparasites. Prior to treatment, animals used several different nest chambers within a single burrow, moving from one chamber to another every 1.6 d on average, and most individuals also slept in more than one burrow. Anti‐parasite treatment reduced the rate at which animals switched from one nest chamber to another within a burrow. Screening of a separate sample of animals for ectoparasites revealed that they were infested with fleas, mainly Xenopsylla eridos. We suggest that by switching periodically from one nest chamber to another, whistling rats reduce the rate at which ectoparasites, especially fleas, accumulate.     

Alteration of burrow‐related behaviour of the norway lobster,nephrops norvegicus during infection by the parasitic dinoflagellate hematodinium

This study has used time‐lapse video recordings to analyze the burrow‐related behaviour of N. norvegicus under a 12 h light: dark regime in aquarium conditions, and compared this with the behaviour of lobsters infected by the parasitic dinoflagellate Hematodinium. Uninfected lobsters performed a higher number of burrow departures, of longer duration, during the hours of darkness than during the light. Infected lobsters performed a relatively constant number of departures of similar‐duration in the light and the dark periods. However, the mean number of departures per day performed by infected lobsters (70 day^‐1) was more than double that of uninfected lobsters (30.1 day^‐1), while the mean duration of burrow departures performed by infected lobsters (258.5s departure^‐1) was more than six times greater than in uninfected lobsters (38.7s ? departure^‐1). This led to a more than ten‐times increase (from 1.7 to 19.4%) in the percentage of the day spent out of the burrow by infected lobsters. This altered burrow‐related behaviour could be due to the nutritional demands of the parasites on infected lobsters, causing an increased requirement to forage, or alternatively to a ‘functional hypoxia’, due to the additional respiratory demands of the parasite, causing the host to emerge from the burrow onto the sediment surface to seek oxygen. Implications for the increased time spent out of the burrow are discussed in relation to availability of infected lobsters by trawlers and predators, and to the use of infection prevalence data from trawl‐caught samples in stock assessment models for N. norvegicus.     

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.     

Trilobites and minute ovoid pellets in a burrow (Ordovician,llanvirnian, Czech Republic)

Old collections from the Ordovician of the Barrandian area (Czech Republic) have yielded a probable vertical burrow that contains several dozen specimens of the pliomerid trilobite Placo‐paria (Placoparia) cambriensis Hicks, some of them enrolled. The fill of the burrow (classified as ?Skolithos isp.) also contains minute ovoid pellets corresponding to the ichnotaxon Tomaculum cf. problematicum Groom. The enrollment of a portion of the trilobite exoskeletons might reflect a defensive response. The pellets may be interpreted as fecal material or, less probably, as eggs. The exoskeletons and the pellets might have been mechanically trapped into the open burrow, or their concentration may have resulted from biological relationships between the tracemaker and the trilobites.     

Development of an index of abundance for pygmy rabbit populations

The pygmy rabbit (Brachylagus idahoensis) is a cryptic, burrowing lagomorph of conservation concern for which an efficient method to monitor populations is needed for conservation planning. We developed an index of abundance based on density of active burrow systems at 7 sites (57.2–118.5 ha) in east central Idaho. We conducted censuses of burrow systems and used mark-resight surveys of 80 radio-collared individuals to estimate density of rabbits. At 5 sites, we also used a second method to estimate rabbit numbers based on presence of tracks in snow around burrow systems. We evaluated patterns of burrow use by individuals and examined the relationship between vegetation structure and density of rabbits. Density of active burrow systems varied from 0.19 to 3.46 per ha, and density of rabbits ranged from 0.02 to 0.46 per ha. Number of burrow systems used by individuals increased with density of available burrows, which supported a nonlinear relationship between abundance of burrows and rabbits. Population density increased curvilinearly with density of active burrows accounting for >75% of the variation (r² = 0.79) in population estimates across sites. We documented a positive relationship between visual obstruction of vegetation and density of rabbits across 6 of the study sites. Our results suggest that density of burrows can serve as an index for monitoring changes in abundance of pygmy rabbits in east central Idaho and that this index also might be useful for monitoring changes in relative abundance over time at other locations. To assess abundance at larger spatial scales or across different regions, the index should be calibrated under regional conditions and site-level covariates should be evaluated. © 2011 The Wildlife Society.     

Application of morphologic burrow interpretations to discern continental burrow architects: Lungfish or crayfish?

A methodology for trace fossil identification using burrowing signatures is tested by evaluating ancient and modern lungfish and crayfish burrows and comparing them to previously undescribed burrows in a stratigraphic interval thought to contain both lungfish and crayfish burrows. Permian burrows that bear skeletal remains of the lungfish Gnathorhiza, from museum collections, were evaluated to identify unique burrow morphologies that could be used to distinguish lungfish from crayfish burrows when fossil remains are absent. The lungfish burrows were evaluated for details of the burrowing mechanism preserved in the burrow morphologies together forming burrowing signatures and were compared to new burrows in the Chinle Formation of western Colorado to test the methodology of using burrow signatures to identify unknown burrows.

Permian lungfish aestivation burrows show simple, nearly vertical, unbranched architectures and relatively smooth surficial morphologies with characteristic quasi‐horizontal striae on the burrow walls and vertical striae on the bulbous terminus. Burrow lengths do not exceed 0.5 m. In contrast, modern and ancient crayfish burrows exhibit simple to highly complex architectures with highly textured surficial morphologies. Burrow lengths may reach 4 to 5 m.

Burrow morphologies unlike those identified in Gnathorhiza aestivation burrows were found in four burrow groups from museum collections. Two of these groups exhibit simple architectures and horizontal striae that were greater in sinuosity and magnitude, respectively. One of these burrows contains the remains of Lysoro‐phus, but the burrow surface reveals no reliable surficial characteristics. It is not clear whether Lysorophus truly burrowed or merely occupied a pre‐existing structure. The other two groups exhibit surficial morphologies similar to those found on modern and ancient crayfish burrows and may provide evidence of freshwater crayfish in the Permian.

Burrows from the Upper Triassic Chinle Formation in western Colorado exhibit simple to moderately complex architectural morphologies, ranging from predominantly vertical, unbranched, with little or no chamber development to predominantly vertical, few branches, and with minor chamber development. Surficial burrow morphologies are moderate to highly textured. The burrows have scrape marks, scratch marks, mud and lag‐liners, knobby surfaces, pleopod striae, and body impressions.

Although no fossil remains of the burrowing organism were found within or associated with the Chinle burrows from western Colorado, the similarity of architectural and surficial burrow morphologies to those in the Chinle of Canyonlands, Utah and to modern crayfish burrows, clearly indicates that the Colorado burrows are the product of burrowing crayfish rather than lungfish. Evaluation of burrowing signatures preserved in the architectural and surficial burrow morphologies is a very useful tool to compare and contrast Chinle burrows from different regions on the Colorado Plateau. Documentation of crayfish burrows in the Chinle of Utah and Colorado strongly suggests that other large‐diameter Chinle burrows elsewhere on the Colorado Plateau and in stratigraphically equivalent units may also be the product of crayfish activity.     

Microscale oxygen distribution in various invertebrate burrow walls

Profiles of dissolved oxygen were measured in pore waters of unburrowed sediment and the burrow walls of seven invertebrate dwellings. Burrows studied include those of Corophium volutator, Heteromastus filiformis, Arenicola marina, Saccoglossus bromophenolosus, Clymenella sp., Hemigrapsus oregonensis and Cirriformia luxuriosa all from mudflats in Willapa Bay, Washington. These animals comprise a range of burrow architectures ranging from simple, unlined burrows to more complex, mucous lined burrows. Oxygen penetrated unburrowed sediment between depths of 0.4–2.6 mm, whereas oxygen penetrated the burrow walls from 0.3 mm to 2.3 mm. Three groups of burrows are recognized based on the oxygen diffusive properties relative to the unburrowed sediment including those that: (1) slightly impeded oxygen penetration, (2) clearly inhibited oxygen penetration, and (3) enhanced oxygen penetration. Differences in the diffusive properties of the burrow wall are related to the burrow microstructure and presumably the microbial communities living within the burrow microenvironment. The results of this study suggest that burrow shape and burrow‐wall architecture may play an important role in controlling the diffusion of oxygen, and possibly of other dissolved gases (i.e. CO₂, H₂S). The results further demonstrate that simplified assumptions (i.e. that bioturbation uniformly enhances oxygen diffusion into suboxic and anoxic sediments), while requisite for numerical modelling, are not necessarily representative of field data.     

Feeling the pressure at home: Predator activity at the burrow entrance of an endangered arid‐zone skink

Habitat modification and invasive species are among the most important contemporary drivers of biodiversity loss. These two threatening processes are often studied independently and few studies have focused on how they interact to influence species declines. Here we assess the predation pressure placed on the threatened great desert skink (Liopholis kintorei) and how this interacts with fire‐induced habitat modifications. We collected daily track data of potential predators for 1 month at 30 great desert skink burrow‐systems where vegetation cover varied significantly after experimental burns. We used these data to evaluate potential predation pressure at the burrow‐system and assess whether fire influenced predator pressure. We supplemented this analysis by documenting predation via the inspection of large mammalian predator scats collected from great desert skink habitat. The level of feral cat activity at a burrow‐system entrance was significantly higher than that of any other potential predator, however fire had no effect on the visitation rates of feral cats, dingoes or large snakes to great desert skink burrow‐systems. The remains of great desert skink were found significantly more frequently in feral cat scats, compared to fox and dingo scats. We provide the first direct evidence that feral cats are a significant predator for great desert skink, thus supporting the hypothesis that feral cat predation is a key threatening process. Feral cat activity was not influenced by small‐scale experimental burns, however, this does not preclude an effect of larger scale fires and we recommend further research exploring this possible interaction.     

Firmground crustacean burrow systems (Glossifungites ichnofacies) in marine shelf deposits,Paleocene Clayton Formation,Alabama, USA

The Paleocene (Danian) Clayton Formation of western Alabama, USA, includes multiple marine shelf parasequences, each comprising a relatively thick marl, capped by a thin limestone, the latter variably reflecting marine flooding episodes. The marls host relatively large firmground burrow systems that penetrate 50–60 cm beneath, and are cast by, superjacent limestones. Excavation of two partially exposed burrow systems – one beneath a highstand parasequence-bounding flooding surface and the other beneath an overlying coplanar sequence boundary/transgressive surface (SB/TS) – reveals complex, primarily horizontal, irregularly branching networks. The former, allied with Thalassinoides paradoxicus, lacks wall bioglyphs, whereas the latter, allied with Spongeliomorpha iberica, is characterized by pervasive, mainly rhombohedral wall bioglyphs that reflect a relatively more firm substrate. Contrasts between these burrow systems are consistent with sequence stratigraphical context and inferred differences in the mechanism and magnitude of depositional hiatuses responsible for firmground development. Both excavated burrow systems likely represent cumulative structures produced by multiple organisms over extended periods of time. The cumulative nature and potential taphonomic biases associated with these and comparable burrow systems in the stratigraphical record preclude confident interpretation of tracemakers and their behaviours. The Clayton burrow systems likely were produced by one or more species of decapod crustacean that engaged in suspension-feeding, surface detritus feeding, gardening or some combination thereof.     

Microvertebrates preserved in mammal burrows from the Holocene of the Argentine Pampas: a taphonomic and paleoecological approach

Microvertebrates are a major component of many assemblages recovered from the Quaternary of the Argentine Pampas. The main goal of this paper is to analyse the taphonomic history of a Holocene microfossil bonebed, recovered from the infilling of a burrow. Evidences suggest the plains vizcacha Lagostomus maximus as the putative producer of the burrow. The assemblage includes individuals belonging to different taxa of mammals (marsupials and rodents) and reptiles (snakes). Taphonomic features suggest that the accumulation inside the burrow was related to flooding processes in the plain. The burrow was a natural trap that favoured the accumulation and preservation of remains corresponding to individuals from different sources. According to the taphonomic evidence, some individuals (Lagostomus maximus, Lestodelphys halli and Serpentes indet.) died inside the burrow, whereas others (Microcavia australis, Reithrodon auritus and Ctenomys sp.) died outside the burrow, and after a time of being exposed on the surface their remains were transported by surface run-offs into the burrow. The record of Lestodelphys halli and Serpentes indet. in the burrow produced by Lagostomus maximus could be related to a circumstantial use. Mammal burrows are a significant taphonomic mode for the late Cenozoic of the Argentine Pampas.     

Rapid escape reflexes in aquatic oligochaetes: variations in design and function of evolutionarily conserved giant fiber systems

Summary This study provides neuroanatomical and electrophysiological evidence that an arrangement of three dorsal giant fibers, functioning as two distinct and dichotomous conduction pathways, has been evolutionarily conserved within the three major orders of aquatic and terrestrial oligochaetes. The medial giant fiber (MGF), activated by afferents of anterior segments, initiates anterior shortening; whereas, the two lateral giant fibers (LGFs), activated in synchrony by afferents of posterior segments, initiate a different response (usually tail withdrawal). Notwithstanding these common features, the design and function of LGF systems differ considerably in aquatic and terrestrial groups. In posterior segments of aquatic species, LGFs are disproportionately larger and conduct faster than MGFs. This contrasts with posterior segments of earthworms in which LGFs are smaller and conduct slower than MGFs.In addition, in aquatic tubificids, a single LGF spike is sufficient to evoke rapid and complete tail withdrawal, whereas a pair of closely-spaced LGF spikes are needed to elicit posterior shortening in earthworms. The graded nature of earthworm escape seems appropriate for worms that burrow in relatively hard substrates and may frequently encounter inanimate stimuli that evoke meaningless giant fiber spiking. On the other hand, the all-or-none nature of the tubificid escape appears advantageous for relatively sedentary worms that are vulnerable to intense predation but reside in aqueous sediments where triggering of giant fiber spikes by non-threatening stimuli is infrequent.Our studies suggest that anatomical and physiological modifications of giant fiber pathways in aquatic and terrestrial worms have occurred during the evolution of oligochaete nervous systems. We hypothesize that differential predation pressures, together with fundamental differences in lifestyle and habitat, have led to this divergence in the structure and function of evolutionarily conserved sets of homologous giant interneurons.Abbreviations HRP horse raddish peroxidase - LGF lateral giant fiber - MGF medial giant fiber - VNC ventral nerve cord     

Burrow architecture,family composition and habitat characteristics of the largest social African mole-rat: the giant mole-rat constructs really giant burrow systems

Among African mole-rats, the giant mole-rat Fukomys mechowii is the largest social species. Despite several attempts to study a free-living population, information on its biology from natural habitats is very scarce. We mapped two neighbouring burrow systems of the giant mole-rat in a miombo woodland in Zambia. We provide information on the size and kin structure of the respective mole-rat families, architecture of their burrow systems, and characteristics of the food supply and soil around the two mapped and additional ten burrow systems. Both uncovered burrow systems were very large (total lengths, 2,245 and 743 m), making them the largest burrow systems ever mapped. Food resources around the additional ten burrow systems had a clumped distribution (standardized Morisita index of dispersion = 0.526), but a relatively high biomass (298 ± 455 g m⁻²). This, together with favourable soil conditions even in the advanced dry season (cone resistance, 328 ± 50 N m⁻²; soil density, 1.36 ± 0.06 g cm⁻³) indicates relatively hospitable ecological conditions. Both food supply and soil conditions were comparable with the conditions found in a miombo habitat of the solitary silvery mole-rat in Malawi. This suggests that there are no ecological constraints which would preclude the solitary life of a subterranean herbivore from the examined habitat. Microsatellite analysis supported the assumption that giant mole-rats live in monogamous multigenerational families with only one breeding pair of non-related animals and their offspring. The mean family size is consistent with previous findings on this species and comparable to that found in other Fukomys species studied thus far.     

On the Ichnotaxonomic Status of Haplotichnus indianensis (Miller, 1889)

The type specimens of the ichnospecies Haplotichnus indianensis and Treptichnus bifurcus were compared under low-angle light. H. indianensis was found to exhibit segments with projections, which are morphological characteristics of T. bifurcus. Consequently, H. indianensis is reinterpreted here as a shallow, subsurface burrow and is considered to be a subjective junior synonym of T. bifurcus. Traces assigned to H. indianensis, other than the type, should be reassigned to other ichnotaxa. Looping surface trails, for example, should be called Gordia isp.     

Ecosystem engineering in a high and cold desert: Effect of a subterranean rodent on lizard abundance and behaviour

Subterranean rodents are often considered as ecosystem engineers because they physically modify the surrounding environment due to their burrowing and foraging activities. Understanding the modifications that ecosystem engineering species exert on the environment are of crucial importance in ecology studies, since they may affect the structure and population dynamics of several species, including lizards. Thus, the objective of the present study is to test the effect that Ctenomys mendocinus exert in the abundance of Liolaemus ruibali and its escape behaviour, in a high-elevation desert. Lizard abundance was estimated using observation transects and escape behaviour was studied with an experiment where the observer was considered by lizards as a potential predator and distance before the lizard flees was measured. All the variables were compared between areas disturbed by C. mendocinus and undisturbed ones. We found that L. ruibali was favoured by C. mendocinus activity. By creating burrow systems that serve as refuges for lizards, this rodent species increases the abundance of L. ruibali and reduces its flight distance, thereby improving its escape performance. We may suggest that C. mendocinus, through the construction of burrow systems, would be acting as an ecosystem engineer in Puna desert, affecting L. ruibali ecology.     

The microenvironment of naked mole‐rat burrows in East Africa

Naked mole‐rats (Heterocephalus glaber) can be extremely long‐lived and are resistant to cancer. Hence, they have been proposed as a model organism for delayed ageing. Adaptation to a constant hypoxic and hypercapnic environment has been suggested as reason for their apparent ability to tolerate oxidative stress. Nevertheless, little is known about the natural habitat to which the species evolved. Naked mole‐rat burrow environments were assessed in Ethiopia and Kenya. Despite reported thermolability of naked mole‐rats, skin temperature upon capture varied (23.7–35.4°C), mostly within the species’ thermoneutral zone, demonstrating their ability to maintain homoiothermy even under wide fluctuations of burrow temperature (24.6–48.8°C) and humidity (31.2%–92.8%), which are far greater than previously reported. Burrow temperature regularly alternates during the daytime and night‐time, driving convective currents that circulate air in the tunnels. Consequently, concentrations of CO₂ and O₂ in burrows only slightly deviated from surface atmosphere. This contradicts the assumption of constant hypoxia/hypercapnia in subterranean burrows. In addition to diffusion, animal movement and occasional wind‐driven ventilation, our data support the temperature‐driven convective model of circulation. The naked mole‐rat burrow is a relatively normoxic subterranean microenvironment with considerable fluctuations in temperature and humidity.     

Effects of estuarine infauna on sediment stability and particle sedimentation

Secretions produced by the two estuarine benthic invertebrates Corophium volutator and Nereis diversicolor stabilise sediments by increasing their shear strength, and reduce sedimentation of previously resuspended particles. The secretions consist of 1 to 2 m threads which bind the particles together. They are used by both species in the production of complex burrow systems in the top 15 cm of sediment. The burrow systems have been demonstrated by a new resin impregnation technique. These results have widespread implications for the stability and erosion of estuarine sediments.     

The effect of particle size frequency distribution of the substratum on the burrowing ability of Chiridota rigida (Semper) (Echinodermata: Holothuroidea)

Chiridota rigida (Semper) burrows easily into well‐sorted substrata, moving between particles in coarse substrata and pushing aside particles in fine substrata. Chiridota rigida does not burrow easily into poorly sorted substrata because spaces which can be penetrated are not available and because the particles cannot be moved as a result of increased stability of the substratum. In poorly sorted substrata, burrowing ability increases with increases in the proportion of fine particles. It would appear that the distribution and abundance of Chiridota rigida and other chiridotids would be affected by the effect of the substratum on their ability to burrow.