Burrowing dynamics and energy cost of transport in the soft-bodied marine invertebrates Polyphysia crassa and Priapulus caudatus

The dynamics and mechanical forces generated during burrowing in Polyphysia crassa (Annelida: Polychaeta) and Priapulus caudatus (Priapulida) were investigated. Both animals live in soft marine muds and burrow by utilizing a direct peristaltic wave alternating with a high internal pressure event which thrusts the anterior part of the body into the substratum. Forces generated during the various phases of a typical burrowing cycle were measured in animals moving beneath the natural substratum at 5±3 °C using electronic transducers and recorder. During 'head' advance Polyphysia generated 0.027 N, and during 'tail' advance 0.020 N, with peak internal pressures averaging 0.95 kPa (= 0.095 N/cm²). Force by Priapulus during head advance and tail advance was 0.081 N and 0.121 N, respectively, with peak internal pressures averaging 2.47 kPa (= 0.247 N/cm²). Polyphysia moves more slowly (0.24 cm/min) than does Priapulus (0.76 cm/min) and expends more energy on mass moved per unit distance. These force measurements during a burrowing cycle were used in place of respirometry as a basis for computation of net cost of transport (NCT) for each animal. NCT for Polyphysia was 635 J kg^-1 m^-1 and for Priapulus was 314Jkg^-1m^-l. Cost of transport for all burrowing animals thus far investigated is high compared to swimming, running and flying. For soft-bodied invertebrates that live an entirely buried existence this high cost must be interpreted in the broader context of the adaptive value of infaunal life, especially protection against predation, and not as simply a means of moving about.     

The burrowing of Nereis diversicolor O.F. Müller,together with some observations on Arenicola marina (L.) (Annelida: Polychaeta)

The mechanism of burrowing in Nereis diversicolor O.F. Müller is described. Proboscis eversion in order to make a hole in the substratum is accompanied by a coelomic pressure pulse of 1.2–7.5 kPa, compared with a value of ≈0.3 kPa during ambulatory activity. Forward movement of the tail in the later stages of burial is by longitudinal contraction, usually accompanied by a pulse of about 2 kPa. The septa appear to act as effective hydrostatic pressure barriers during burrowing.In the flange-proboscis sequences of Arenicola marina (L.) the coelomic pressure in the trunk during erection of the flanges is generally slightly higher than that in the head, while the reverse applies during the low-volume proboscis eversion. Major progression of the anterior usually occurs during the first flange-proboscis sequence of each cycle and that of the posterior early in the dilation sequence. External pressure recordings of irrigation-defaecation cycles are given.Nereis uses a similar method of burrowing to that of the free-moving Nephtys but the coelomic pulses of the former are lower, as in the rate of burial (about 0.02 cm s^?1) which is similar to that of Arenicola. Closure of the septa of Nereis during burial suggests that its burrowing mechanism is not as versatile as that of Nephtys.     

Image analysis of the burrowing mechanisms of Polyphysia crassa (Annelida: Polychaeta) and Priapulus cudatus (Priapulida)

The burrowing mechanisms of two soft-bodied marine invertebrates, Polyphysia crassa (Annelida: Polychaeta) and Priapulus caudatus (Priapulida) were re-examined and extended utilizing computer image analysis. Predetermined points on sequential photographs of animals burrowing in methylcellulose (an artificial medium of high transparency) were digitized and stored as x, y coordinates from which were calculated segment length, width, and volume in addition to producing real and straightened images of the animal on a plotter. Both species are adapted to soft substrata and both utilize a direct peristaltic wave to advance the body into a cavity or loosened area formed by anterior structures. Polyphysia displaces and loosens the substratum with lateral scraping movements of its anterior six segments (the "head region") while Priapulus makes a large cavity anteriorly by the forceful eversion and dilation of the praesoma. Unlike Priapulus, Polyphysia utilizes a different method of locomotion when moving on the mud surface as opposed to burrowing, the former involving two direct peristaltic waves at a time and negligible internal pressures resulting in nearly continuous advance. However when burrowing, Polyphysia , like Priapulus , utilizes a single direct peristaltic wave alternating with phasic pressure pulses which advance the animal in step-wise fashion.     

Evidences of habitat displacement between two common soft-bottom SW Atlantic intertidal crabs

The intertidal burrowing crab Chasmagnathus granulatus Dana is the dominant species in soft sediments and vegetated intertidal areas along the SW Atlantic estuaries (southern Brazil 28°S to the northern Argentinean Patagonia 41°S) where it produces dense and extensive burrowing beds. The mud crab Cyrtograpsus angulatus Dana coexists with Ch. granulatus in this area, but it also inhabits areas to the south (northern and central Argentinean Patagonia). A survey covering both areas showed that C. angulatus rarely live in burrows when coexisting with Ch. granulatus, but form large burrowing beds when not coexisting with Ch. granulatus. When both species coexisted, burrowing beds of C. angulatus are restricted to sandy-muddy areas. Only rarely are burrows of C. angulatus found within Ch. granulatus beds. However, when Ch. granulatus were experimentally excluded within their burrowing beds, new settlers of C. angulatus made burrows and maintained them until they reached large size. Paired (inside and outside Ch. granulatus burrowing bed) sampling during high tide using beach nets showed that C. angulatus rarely venture inside the Ch. granulatus crab beds. Other field experiments showed that adults Ch. granulatus always displace C. angulatus from burrows. Furthermore, in several sites located south of the limit of distribution of Ch. granulatus at the Patagonian coast, soft bare intertidals are dominated by burrowing beds of C. angulatus mixed with the congener C. altimanus Dana. Together, these evidences suggest that the mud crab C. angulatus is displaced from soft bottom areas by the burrowing crab Ch. granulatus. It is an example of competitive exclusion through aggressive interference in soft-bottom habitats when the shared resource is the access to sediment surface, a two-dimensional well-defined resource.     

The role of the coelom as a hydrostatic skeleton in lingulid brachiopods

Anatomical and experimental studies of the perivisceral coelom and pedicel of Lingula ana lina indicate that the coelomic fluid functions as a hydrostatic skeleton in respect of valve and pedicel movements, valve opening always being associated with positive pressures. The perivisceral coelom is surrounded by a body wall containing circumferential muscle fibres, whilst all muscles passing between the valves (principally adductor and oblique fibres) are located within the body wall. These two sets of muscles function similarly to the circular and longitudinal muscles of a classical hydrostatic skeleton.
Pressure recordings from the lumen of the pedicel and perivisceral coelom, during opening or rotary movements of the valves, were similar and showed pressure pulses of up to 0.8 kPa. During the initial stages of burrowing, pulses of up to 2.5 kPa were observed when the valves were being pressed into the sand. These values are well within the capability of the circumferential muscles of the body wall.     

Cuticle expansion during feeding in the tick Amblyomma hebraeum (Acari: Ixodidae): The role of hydrostatic pressure

Female Amblyomma hebraeum ticks (Acari: Ixodidae) increase their weight ∼10-fold during a ‘slow phase of engorgement’ (7–9 days), and a further 10-fold during the ‘rapid phase’ (12–24 h). During the rapid phase, the cuticle thins by half, with a plastic (permanent) deformation of greater than 40% in two orthogonal directions. A stress of 2.5 MPa or higher is required to achieve this degree of deformation (Flynn and Kaufman, 2015). Using a dimensional analysis of the tick body and applying the Laplace equation, we calculated that the tick must achieve high internal hydrostatic pressures in order to engorge fully: greater than 55 kPa at a fed:unfed mass ratio of ∼20:1, when cuticle thinning commences (Flynn and Kaufman, 2011). In this study we used a telemetric pressure transducer system to measure the internal hydrostatic pressure of ticks during feeding. Sustained periods of irregular high frequency (>20 Hz) pulsatile bursts of high pressure (>55 kPa) were observed in two ticks: they had been cannulated just prior to the rapid phase of engorgement, and given access to a host rabbit for completion of the feeding cycle. The pattern of periods of high pressure generation varied over the feeding cycle and between the two specimens. We believe that these pressures exceed those reported so far for any other animal.     

Burrowing behavior of Dermatonotus muelleri (Anura,Microhylidae) with reference to the origin of the burrowing behavior of Anura

Dermatonotus muelleri is a forelimbs-head-first burrowing frog that uses its forelimbs for soil removal, and it is the second anuran species known to arch its head downwards at an angle of almost 90° to the longitudinal axis of its body when burrowing. The burrowing behavior of D. muelleri is divided in three stages: head burrowing, body burrowing, and chamber construction. Burrowing in D. muelleri includes construction of a subterranean chamber used for estivation during the dry season. Phylogenetic analysis based on literature survey of burrowing behavior suggested that head-first burrowing behavior has evolved several times in anuran history, forming a convergence complex, and that hindlimbs-first burrowing is a basal behavior.     

Repeatability of subject/bed interface pressure measurements

The repeatability of a technique for measuring interface pressures has been assessed. Pressure was measured using a Talley SA500 Pressure Evaluator under six anatomical sites (occiput, scapula, elbow, sacrum, buttock, and heel) of six healthy subjects lying supine on a Clinifloat mattress (trademark, SSI). For each site of each subject, four repeat readings were taken per day on four separate days. Mean pressures varied significantly between subjects (p < 0.02), though differences in mean pressures between sites were greater. Pressure was not significantly related to subject mass. The overall repeatability of the technique was ± 0.77 kPa (± 5.8 mm Hg) which was much smaller than the range of pressures found under different sites (2.72 kPa or 20.4 mm Hg at the sacrum to 9.00 kPa or 67.5 mm Hg at the heel). Repeatability varied from site to site, from ± 0.47 kPa (± 3.5 mm Hg) at the buttocks to ± 1.20 kPa (± 9.0 mm Hg) at the heel. Measurements were found to vary significantly more between days than between repeats on the same day (p < 0.02).     

Establishing predictive indicators for the status of loaded soft tissues.

Two complementary techniques were employed to assess the soft tissue response to applied pressure. The noninvasive methods involve the simultaneous measurement of the local tensions of oxygen and carbon dioxide (tcPO2 and tcPCO2) and the collection and subsequent analysis of sweat collected from the sacrum, a common site for the development of pressure sores. All tests were performed on able-bodied subjects. Results have indicated that oxygen levels (tcPO2) were lowered in soft tissues subjected to applied pressures of between 40 (5.3 kPa) and 120 mmHg (16.0 kPa). At the higher pressures, this decrease was generally associated with an increase in carbon dioxide levels (tcPCO2) well above the normal basal levels of 45 mmHg (6 kPa). There were also considerable increases, in some cases up to twofold, in the concentrations of both sweat lactate and urea at the loaded site compared with the unloaded control. By comparing selected parameters, a threshold value for loaded tcPO2 was identified, representing a reduction of ~60% from unloaded values. Above this threshold, there was a significant relationship between this parameter and the loaded/unloaded concentration ratios for both sweat metabolites. These parameters may prove useful in identifying those subjects whose soft tissue may be compromised during periods of pressure ischemia.     

Disturbance,emigration, and refugia: How the mud snail,Ilyanassa obsoleta (Say), affects the habitat distribution of an epifaunal amphipod,Microdeutopus gryllotalpa (Costa)

In the presence of the mud snail Ilyanassa obsoleta (Say), the tubicolous amphipod Microdeutopus gryllotalpa (Costa) emigrates to snail-free sediments, as demonstrated in laboratory and field experiments. Emigration occurs predominantly in the dark when the amphipod is most active. Unlike crevices, the thickness of sediments in which the amphipod is established offers no protection from snail disturbance. Emigration is shown to be caused by the disturbance generated by the snail's plowing and burrowing across the sediment surface, and not a response to a reduction in the shared microfloral food supply. The crawling and burrowing of the smaller mud snail, Hydrobia totteni Morrison, does not disturb Microdeutopus, supporting the hypothesis that relative body sizes affects the ability of bioturbators/burrowers to disturb tube-dwellers. As the burrows of Microdeutopus extend only ≈2 cm below the sediment surface, thick mud layers do not offer any refuge from Ilyanassa. However, very small solid surfaces (≈1–2 mm in relief) to which the amphipods build tubes do provide some protection from Ilyanassa. In soft-sediment benthic communities, such small structures may provide significant refuge for small epifauna and shallow burrowing infauna escaping from small-scale, biogenic disturbance.     

Interactions between locomotion,feeding, and bodily elongation during the evolution of snakes

Information from lizard lineages that have evolved a highly elongate (snake‐like) body form may clarify the selective forces important in the early evolution of snakes. Lizards have evolved bodily elongation via two distinct routes: as an adaptation to burrowing underground or to rapid locomotion above ground. These two routes involve diametrically opposite modifications to the body plan. Burrowing lizards have elongate trunks, small heads, short tails, and relatively constant body widths, whereas surface‐active taxa typically have shorter trunks, wider heads, longer tails, and more variable body widths. Snakes resemble burrowing rather than surface‐active (or aquatic) lizards in these respects, suggesting that snakes evolved from burrowing lizards. The trunk elongation of burrowing lizards increases the volume of the alimentary tract, so that an ability to ingest large meals (albeit consisting of small individual prey items) was present in the earliest snakes. Subsequent shifts to ingestion of wide‐bodied prey came later, after selection dismantled other gape‐constraining morphological attributes, some of which may also have arisen as adaptations to burrowing through hard soil (e.g. relatively small heads, rigid skulls). Adaptations of snake skulls to facilitate ingestion of large prey have evolved to compensate for the reduction of relative head size accompanying bodily elongation; relative to predator body mass, maximum sizes of prey taken by snakes may not be much larger than those of many lizards. This adaptive scenario suggests novel functional links between traits, and a series of testable predictions about the relationships between squamate morphology, habitat, and trophic ecology. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 293–304.     

Locomotion in Nebalia bipes: a possible model for Palaeozoic phyllocarid crustaceans

The morphofunctional aspects of locomotion in Recent phyllocarid crustaceans are presented, based on combined video observations of living specimens of the cosmopolitan nektobenthic Nebalia (Leptostraca, Nebaliidae) and scanning electron microscopy of fixed material. N. bipes is infaunal by day, showing a preference for organic-rich black muds and dim light conditions. Under natural conditions, emergence from sediment and nocturnal swimming activities are controlled by a circadian rhythm. The 1st and 2nd antennae perform the major role in digging. The carapace rostral plate acts as a ram diverting particles and preventing inputs of sediment into the carapace. Superficial burrowing may provide camouflaged shelter against predation. Swimming is accomplished by the combined action of the first four pairs of pleopods (backward metachronal flexion during the power stroke) and the well-articulated abdominal shaft (downward flap). Plumose setae present on both the pleopods and furcal rami (governed by low Reynolds numbers) behave like paddles maximizing the resistance to the water. Clusters of cuticular microscales (3-5 μm) and microdenticles are present on the external surface of the carapace and the trunk segments, respectively. Both show a uniform directional arrangement suggesting functional interpretations in relation to locomotion (e.g., to prevent back slippage during digging, to reduce turbulence in the flow layer close to the body, and/or to serve as mechanoreceptors for the detection of variation in water flow). Key features of functional importance in the locomotion of living leptostracans (natatory pleopods, a highly flexible abdomen, setulate or paddle-like furcal rami) are also recognized in Palaeozoic archaeostracan phyllocarids, suggesting that swimming was accomplished by the same pleopodal movements as described for N. bipes. Comparisons with Recent analogues (Nebaliopsis) suggest that phyllocarids with a free swimming life-style existed among the lower Palaeozoic archaeostracans (e.g., Caryocaris; Ordovician). Asymmetrical cuticular ornament in some archaeostracans indicates burrowing habits. Some Palaeozoic phyllocarids may have been occasional (possibly diurnal) mud dwellers comparable to modern nebaliids.     

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 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.     

SEASONAL OCCURRENCE OF XYLEM EMBOLISM IN SUGAR MAPLE (ACER SACCHARUM)

Xylem embolism, the reduction of water flow by air-filled vessels, was measured in a stand of 5- to 8-year-old sugar maple (Acer saccharum Marsh.) saplings growing in a nursery bed in northwestern Vermont. Embolism was quantified as percentage loss in hydraulic conductivity of trunk and branch segments relative to maximum values obtained by removing air from vessels by repeated high pressure (173 kPa) perfusions. Ten segments per tree were cut from 6 trees for each of 11 measurement periods spaced at roughly monthly intervals from May 1986 to June 1987. During the 1986 growing season, embolism increased significantly from 11 to 31% in the larger branches and trunk (segment diameter #8805;0.5 cm), but remained at about 10% in twigs (segment diameter <0.5 cm). This was unexpected because the greatest water stess and thus potential for embolism occurs in twigs. During the winter, embolism increased throughout the trees and the trend with diameter was reversed; by February, small twigs were 84% embolized vs. 69% for larger branches and trunk. Dye perfusions showed that winter embolism in trunks was localized on the south side; this may have resulted from water loss by sublimation or evaporation in the absence of water uptake. Beginning in late March, embolism decreased throughout the trees to approximately 20% in June. This decrease was associated with positive xylem pressure of at least 16 kPa which may have originated in the roots, because weather conditions at the time were unfavorable for the generation of stem pressures characteristic of Acer species in early spring.     

The effects of burrowing of Helice tridens (De Haan) on the soil of a salt-marsh habitat

The physical and chemical effects of the burrowing activity of the mud crab Helice tridens (De Haan) on the soil of a salt-marsh habitat were investigated. Soil-turnover rate caused by burrowing activity was found to be ≈ 3% of the soil from the surface to a depth of 40 cm every day during the summer. The vertical distributions of leaf and stem fragments of the salt-marsh plant Phragmites australis (Trin.) and the vertical distribution of ammonium N concentration in the soil were also investigated. At locations in the marsh where there were many large burrows, numerous leaf and stem fragments were recognized in the soil, while in areas in the marsh containing only a few small burrows these fragments were scanty. The soil depths at which leaf and stem fragments were abundant, corresponded to the depths of the burrows. These results show that mud crabs bury fallen plant fragments in the soil by their burrowing activity. Ammonium N in the soil was also abundant at locations in the marsh where there were many burrows, indicating that organic matter, such as fallen leaves and stems, may be decomposed to inorganic nutrients which are useful to the salt-marsh plants.     

Gender differences in regional body composition and somatotrophic influences of IGF-I and leptin.

This study evaluated the arm, trunk, and leg for fat mass, lean soft tissue mass, and bone mineral content (BMC) assessed via dual-energy X-ray absorptiometry in a group of age-matched (approximately 29 yr) men (n = 57) and women (n = 63) and determined their relationship to insulin-like growth factor I (IGF-I) and leptin. After analysis of covariance adjustment to control for differences in body mass between genders, the differences that persisted (P < or = 0.05) were for lean soft tissue mass of the arm (men: 7.1 kg vs. women: 6.4 kg) and fat mass of the leg (men: 5.3 kg vs. women: 6.8 kg). Men and women had similar (P > or = 0.05) values for fat mass of the arms and trunk and lean soft tissue mass of the legs and trunk. Serum IGF-I and insulin-like growth factor binding protein-3 correlated (P < or = 0.05) with all measures of BMC (r values ranged from 0.31 to 0.39) and some measures of lean soft tissue mass for women (r = 0.30) but not men. Leptin correlated (P < or = 0.05) similarly for measures of fat mass for both genders (r values ranging from 0.74 to 0.85) and for lean soft tissue mass of the trunk (r = 0.40) and total body (r = 0.32) for men and for the arms in women (r = 0.56). These data demonstrate that 1) the main phenotypic gender differences in body composition are that men have more of their muscle mass in their arms and women have more of their fat mass in their legs and 2) gender differences exist in the relationship between somatotrophic hormones and lean soft tissue mass.     

Assessment of mechanical conditions in sub-dermal tissues during sitting: a combined experimental-MRI and finite element approach

A common but potentially severe malady afflicting permanent wheelchair users is pressure sores caused by elevated soft tissue strains and stresses over a critical prolonged period of time. Presently, there is paucity of information regarding deep soft tissue strains and stresses in the buttocks of humans during sitting. Strain and stress distributions in deep muscle and fat tissues were therefore calculated in six healthy subjects during sitting, in a double-donut Open-MR system, using a "reverse engineering" approach. Specifically, finite element (FE) models of the undeformed buttock were built for each subject using MR images taken at the coronal plane in a non-weight-bearing sitting posture. Using a second MR image taken from each subject during weight-bearing sitting we characterized the ischial tuberosity sagging toward the sitting surface in weight-bearing, and used these data as displacement boundary conditions for the FE models. These subject-specific FE analyses showed that maximal tissue strains and stresses occur in the gluteal muscles, not in fat or at the skin near the body-seat interface. Peak principal compressive strain and stress in the gluteus muscle were 74+/-7% and 32+/-9 kPa (mean+/-standard deviation), respectively. Peak principal compressive strain and stress in enveloping fat tissue were 46+/-7% and 18+/-4 kPa, respectively. Models were validated by comparing measured peak interface pressures under the ischial tuberosities (17+/-4 kPa) with those calculated by means of FE (18+/-3 kPa), for each subject. This is the first study to quantify sub-dermal tissue strain and stress distributions in sitting humans, in vivo. These data are essential for understanding the aetiology of pressure sores, particularly those that were recently termed "deep tissue injury" at the US National Pressure Ulcer Advisory Panel (NPUAP) 2005 Consensus Conference.     

Habitat preferences of two estuarine burrowing crabs Helice crassa Dana (Grapsidae) and Macrophthalmus hirtipes (Jacquinot) (Ocypodidae)

The horizontal and vertical distributions of two species of endemic, burrowing mud crabs Helice crassa Dana 1851 (Grapsidae) and Macrophthalmus hirtipes (Jacquinot 1853) (Ocypodidae) are described for the Avon-Heathcote Estuary (43°33′S: 172°44′E), Christchurch, New Zealand. Substratum preference is shown to be the most important factor influencing mud crab distribution, but lack of tolerance to salinities below 4‰ is also a significant factor preventing M. hirtipes from occurring at points close to freshwater input. Both species had similar sediment organic content and particle size requirements. Helice crassa was concentrated in well-drained, compacted sediments above mid-tide level, whilst Macrophthalmus hirtipeswas found in waterlogged areas below mid-tide level. This vertical separation is shown not to be caused by differential desiccation tolerances, but by feeding and burrowing adaptations related to these different substrata.     

Density and distribution in laboratory populations of midge larvae (Chironomidae: diptera)

The troglobitic amphipod crustacean Crangonyx antennatus occupies mud-bottom pools and small, gravel-bottom streams in caves in the southern Appalachians. One large, mud-bottom pool population in Lee Co., Virginia was observed periodically from 1967 to 1975. Amphipods in this population frequently burrowed into the soft mud substrate, where they were able to survive desiccation during periods when the pool dried up. Animals kept in the laboratory also burrowed and survived desiccation during an experiment which simulated drought conditions similar to those observed in caves. Amphipods collected from both pool and stream habitats burrowed, thereby indicating that stream-adapted populations of this species still retain sufficient flexibility to survive under variable environmental conditions that might be encountered in nature. It is concluded that burrowing provides a means of survival for C. antennatus when it is sometimes exposed to drought conditions in cave pool habitats and also provides protection from potential terrestrial predators under similar conditions. Burrowing also offers amphipods protection from aquatic predators (such as salamander larvae) during normal water levels and possibly allows juveniles a means of escaping cannibalism by adults.