Activity periods and energetic reserves of the brown lemming in northern Alaska

Radiotelemetry provided data on the activity periods of brown lemmings Lemmus sibiricus in northern Alaska. Activity inside the burrow and inactivity outside the burrow usually occurred in short episodes (<3 min), whereas periods of activity outside the burrow and inactivity inside the burrow were more prolonged and appeared to be associated with foraging (outside) and sleeping (inside). Energetic reserves for lemmings entering a burrow consisted of body fat and undigested gut contents, which allowed minimal survival times of 8.6 ± 2.3 h (mean ± 1 S.E.). Nevertheless, lemmings usually left their burrows within 40 min of entering, at which time their stomachs would be at least two-thirds empty. Activity patterns of lemmings appear to change in response to predators, weather and nutritional requirements. We argue that lemmings seek to minimize the time spent foraging rather than to maximize the net energy gain while foraging.     

The influence of distance to burrow on flight initiation distance in the woodchuck, Marmota monax

We used woodchucks (Marmota monax) to test predictions of acost-benefit model of antipredator behavior that flight initiationdistance would increase with distance to refuge and with predatorapproach velocity. We also examined the effects of distanceto refuge and predator approach velocity on escape velocityand on both temporal and spatial margin of safety (expectedtime and distance between predator and burrow at the time ofthe woodchuck's arrival). The observer, assumed to be perceivedas a potential predator, approached juvenile woodchucks fromthe direction opposite to the burrow at a slow (1.24 m/s) orfast (1.79 m/s) walking pace. When the woodchuck started toflee, the observer recorded the woodchuck's distance from theobserver and from its burrow, the time spent running, and whetherthe woodchuck stopped before reaching its burrow. Flight initiationdistance increased consistendy with distance to the burrow overthe entire observed range (0–25 m) but was not significantlyaffected by observer approach velocity. Escape velocity wasnot significantly influenced by the observer approach velocityand was approximately constant over the range of 2–25m, but was slower for woodchucks less than 2 m from their burrows.Both temporal and spatial margins of safety increased with distancefrom the burrow. The temporal margin of safety increased withdistance from the burrow more rapidly for slow than for fastobserver approach velocity. Woodchucks fleeing from greaterthan 2 m usually stopped near the burrow before entering, butthose from closer distances usually entered directly. Theseresults support the assumption that antipredator behavior issensitive to the costs and benefits of alternative escape decisions.     

The behaviour of Corophium volutator (Crustacea: Amphipoda)

The behaviour of Corophium volutator (Pallas) is outlined. Swimming, crawling, burrowing and feeding activities are described in detail. Animals usually swim on their backs. Every few seconds, swimming alternates with passive sinking. Animals can crawl over surfaces in and out of water. Out of water they do so by a looping motion using their second antennae and telson. When out of water animals crawl away from light and down slopes. In water they swim towards light. Burrowing is initiated by rapid beating of the pleopods; the animal then sinks below the surface by a concerted action of pereiopods, pleopods, telson, uropods and second antennae; within a few minutes, a shallow burrow is formed. The formation of permanent burrows is dependant on particle size, on adhesive properties of detritus and primary films on sand particles, and on a secretion produced by the animal itself. Individuals can turn about in permanent burrows. The species is essentially a detritus feeder. Animals normally feed only when in their burrows, by using their second antennae to scrape material from the substrate surface into the entrance of the burrow. This material is then transported to the mouth by the feeding appendages and respiratory current. The behaviour of small and large animals differs; small animals burrow rapidly and permanently, and do not emerge spontaneously; furthermore, they only swim occasionally. Large animals swim more frequently, spend more time on the substrate surface, and periodically move burrows. It is suggested that new habitats are colonized by large animals which have already bred once.     

Tool Use by Spiders: Stone Selection and Placement by Corolla Spiders Ariadna (Segestriidae) of the Namib Desert

The corolla spider (Araneae: Segestriidae: Ariadna) of the Namib Desert gravel plains typically places seven or eight stones in a circle around its burrow entrance. It was examined whether the spider selects and places stones according to physical characteristics. Circle composition can be explained by the allometric scaling to spider size of the burrow entrance and the stones. The stones were usually placed with the narrowest side or point towards the burrow. Quartz crystals were preferred to four other stone types available. The spiders could detect prey brushing the outside of the circle stones that were about as wide as the spider's path of attack. I conclude that corolla spiders use the selected stones as tools to extend their foraging range.     

Behavioural and physiological adaptations to a burrowing lifestyle in the snake blenny, Lumpenus lampretaeformis, and the red band-fish, Cepola rubescens

Observations have been made on the mode of burrow construction in the snake blenny, Lumpenus lampretaeformis , under laboratory conditions. It appears that head probing and lateral oscillations of the body are principally responsible for the excavation of the burrow which is completed within 24 h. The burrow structure has been analysed in detail, showing a mean depth of 7.2 cm with a maximum observed length of 73 cm, with most systems between 20 and 35 cm in length. Initially linear burrows with two openings are usually provided with a small side tunnel, giving the system a characteristic Y-shape.
Burrow irrigation was investigated for the first time in L. lampretaeformis. The mean duration of burrow irrigation, by flexions of the tail of the fish, was 21 s with over 13 min h⁻¹ spent in irrigating the burrow. The mean water displacement per irrigation period was 3.1 ml. The PO ₂ and PCO ₂ were measured in both surface water and within the burrow system of L. lampretaeformis. Surface water values for PO ₂ were high (> 150 Torr) and PCO ₂ low (<0.4 Torr). Hypoxic and hypercapnic conditions were measured in the burrow system itself, with PO ₂ values ranging between 57 and 129 Torr and PCO ₂ rising to > 1.3 Torr in some burrows.
A comparative study of Cepola rubescens burrows indicated similar surface water PO ₂ and PCO ₂ values as in L. lampretaeformis. Burrow water PO ₂ values ranged between 60 and 94 Torr, with PCO ₂ values as high as 1.5 Torr being recorded. These results are discussed in relation to the adaptation of both species to a burrowing lifestyle.     

Foraging behavior of golden hamsters (<Emphasis Type="Italic">Mesocricetus auratus</Emphasis>) in the wild

Foraging theory posits that animals should maximize energy gains while minimizing risks, the largest of which is usually predation. For small burrowing mammals the best measure of risk avoidance may be the time spent in the burrow, although this measure is rarely examined. During the spring of 2005 and 2006 we recorded the foraging behavior of female golden hamsters in their natural habitat in southern Turkey. Data were collected with a data logger and by direct observations. Female golden hamsters averaged 64 min per day above ground in a series of foraging trips with a mean duration of 5.5 min. Two nursing females increased their time out of the burrow by a factor of 6–8 times over the course of 16 days by increasing both the number of trips and the length of each trip. These results show that hamsters spend little time out of the burrow, thus minimizing risk, but they also show that time spent out of the burrow is related to the energy needs of the hamsters; lactating females with high energy needs exposed themselves to much greater risk than did non-lactating females.     

The role of mounds in promoting water-exchange in the egg-tending burrows of monogamous goby, Valenciennea longipinnis (Lay et Bennett)

Valenciennea longipinnis spawns monogamously in a burrow. After spawning, the paired female constructs a conspicuous mound on the burrow by carrying and piling up substratum-derived materials while the male tends eggs in the burrow until hatching occurs. In this study, the mounds of V. longipinnis were tested in the field to confirm their function of promoting water-exchange in the burrow, and their ecological role was examined in relation to egg care by the male. The mound of V.longipinnis promoted water-exchange in the burrow, contributing to the provision of external oxygenated sea water into the burrow. Therefore, dissolved oxygen (DO) concentrations in the burrow with a mound were significantly higher than those without a mound. Although male egg-tending behaviour (e.g., fanning) may also promote water-exchange in the burrow, the water-exchange appeared to depend mainly on the hydrodynamic effect. Removals of the mound and paired female on the day of spawning led to high rates of egg-desertion by males. Since the frequency and time of fanning increase with a decrease of DO concentration in the burrow, the egg-desertion may result from an increased parental cost to males due to the decrease of water-exchange without a mound. This was supported by the fact that the DO concentrations on the day after mound removal were significantly lower in the egg-deserted burrows (measured before desertions) than in burrows not deserted by the male. Moreover, removals of paired females only also led to higher desertion rates. After removal of the female, the mound gradually collapsed by wave action and other factors, and the surface of the mound was covered with planktonic materials. Such a mound of poor quality may provide little water-exchange, which may lead to the egg-desertion by males. These results indicated that mound maintenance by females during the egg-tending period has an important role in the success of parental care by males.     

Are ghost crabs (Ocypode spp.) smaller on human-disturbed sandy beaches? A global analysis

Hydrobiologia - Testing ghost crabs as indicator species usually fail to depict the mechanisms behind low burrow densities in urban coasts, neglecting individual traits as diagnostic variables. We...     

A Comparative Study of Bivalves Which Bore Mainly by Mechanical Means

This account of the boring mechanisms of those bivalve groupswhich bore mainly by mechanical means attempts to show partlyby reference to published accounts of boring and partly fromour own recent observations of certain characteristics of theboring process in the Pholadidae and Petricolidae, that in contrastto the movements of burrowing forms from which originally allthe boring movements derive, the process of boring makes fewdemands on the hydrodynamic system of the bivalve. The characteristicsof the boring process are closely related to the movements inmodern forms having epifaunal or infaunal habits, supportingthe suggestions of Yonge (1963) concerning the origin of thishabit in the Bivalvia. In all groups in which boringis mechanical,the shell forms the boring tool. However, in those groups inwhich boring has its origin in the epifaunal habit, the majorforce applied to the shell in abrading the burrow isprovidedby contractions of the pedal or byssal retractor muscles. Inthe Adesmacea alone, where boring has been derived from a deepburrowing habit, the adductor muscles provide the major forcein abrasion, and the basic digging cycle has become specializedby the addition of the rocking action of the valves which succeedsretraction. In the former group the ligament is retained andprovides the strong outward force with which the shell is heldagainst the wall of the burrow. In the latter group, the ligamentis reduced, allowing the valves to rock, but here the reciprocalaction of the adductors allows the valves to diverge anteriorlyas the large posterior retractor muscle contracts. In the morespecialized species, water pressure plays a minor role, themaximum pressures recorded being associated with actions subordinateto those involved primarily in abrasion, such as rotation inthe burrow or expulsion of debris from the burrow aspseudofeces.The least specialized borers, such as Petricola, resemble burrowingforms in the importance of the hydrodynamic role of the bodyfluids. In all groups there is a tendency for hypertrophy totake place in the muscles which produce the main boring effect,and for their action to be applied with maximum mechanical advantageagainst a fulcrum provided in most cases by the foot.     

The occupation of artificial burrows by Helice tridens (De Haan) and the length of its stay

The mud-crab Helice tridens (De Haan) influences the cycling of matter in salt-marsh ecosystems through its burrowing activity. If the crabs occupy and stay in their burrows for a short time, their burrowing activity will be great, since they will continuously construct new burrows. Therefore, investigation of the relation between the crabs and their burrows is considered to be important. In the present study, the relation between pipes as a form of artificial burrow and their occupation by the crab was analyzed. A close relationship was recognized between the diameter of the pipe opening and the carapace width of the crab which occupied the pipe, while pipes with a length shorter than the depth of the crab burrows were hardly occupied. These results indicate that the diameter and length of an artificial burrow affects the likelihood of its occupation by this species of crab. The length of the crab's stay in this type of artificial burrow was generally 1 day. This result may be related to the field observation that newly made burrows frequently collapse due to water current occurring through tidal action after the crabs have left.     

The Contribution of Crab Burrow Excavation to Carbon Availability in Surficial Salt-marsh Sediments

Geomorphology, vegetation and tidal fluxes are usually identified as the factors introducing variation in the flushing of particulate organic matter (POM) from tidal marshes to adjacent waters. Such variables may, however, be insufficient to explain export characteristics in marshes inhabited by ecosystem engineers that can alter the quantity and quality of POM on the marsh surface that is subject to tidal flushing. In this study we evaluated the balance between transfer of buried sedimentary organic carbon (C) to the marsh surface due to crab excavation (measured from the mounds of sediment excavated from burrows) and outputs of C from the surface due to sediment deposition within crab burrows (estimated from sediment deposited within PVC burrow mimics), in a Southwestern Atlantic salt marsh supporting dense (approximately 70 ind m⁻²) populations of the crab Chasmagnathus granulatus. C excavation by crabs was much greater than deposition of C within crab burrow mimics. Per area unit estimates of the balance between these two processes indicated that crabs excavated 5.98 g m⁻² d⁻¹ and 4.80 mg m⁻² d⁻¹ of total and readily (10 d) labile C, respectively. However, sediments excavated by crabs showed a significantly lower content of both total and readily-labile C than sediment collected in burrow mimics. This indicates that ecosystem engineering by burrowing crabs causes a net decrease in the concentration of C in the superficial sediment layers and, thus, an overall decrease in the amount of C that can be washed out of the marsh by tidal action. Incorporating the in situ activities of ecosystem engineers in models of marsh export should enhance understanding of the function of marshes in estuarine ecosystems.     

Mole activity in woodlands, fens and other habitats

Monks Wood, a deciduous wood of 155 hectares on clay soil, has an estimated population of 400 moles. There is an anastomosing system of permanent burrows, and when a mole is trapped its area of burrow is soon occupied by another animal. Little new digging occurs, except in winter, after frost has driven the soil fauna deeper into the ground. The burrows usually act as pit-fall traps for the food. Young animals migrating in summer may live more superficially, in tunnels in moss and grass, but survivers usually move into a permanent tunnel system by autumn.
Woodwalton Fen has an easily worked peat soil, and few earthworms. Here moles burrow at all seasons, presumably needing an extensive burrow system to catch sufficient food. In times of flood, the moles leave submerging ground but return very soon after the water subsides. They probably swim across flooded areas but iis yet we do not know if the same animals return after floods to the same burrows.
Moles are not good indicators of soil fertility, particularly as many mole heaps are made by few moles in poor soil, and fewmew heaps may appear in good soil with permanent burrows. We do not understand why there are so few moles in some apparently suitable pasture, with high worm populations, or how others manage to obtain sutficient food in infertile areas.     

Burrow ventilation and associated porewater irrigation by the polychaete Marenzelleria viridis

Burrow ventilation of benthic infauna generates water currents that irrigate the interstices of the sediments surrounding the burrow walls. Such activities have associated effects on biogeochemical processes affecting ultimately important ecosystem processes. In this study, the ventilation and irrigation behavior of Marenzelleria viridis, an invasive polychaete species in Europe, was analyzed using different approaches. M. viridis showed to perform two types of ventilation: (1) muscular pumping of water out of the burrow and (2) cilia pumping of water into the burrow. Flowmeter measurements presented muscular pumping in time averaged rates of 0.15 ml min⁻¹. Oxygen needle electrodes positioned above the burrow openings revealed that muscular undulation of the worm body pumps anoxic water out of the burrow. On the other hand, microscope observations of the animal showed that ventilation of oxygen-rich water in the burrow occurs by ciliary action. The volume of water irrigated by M. viridis appears to vary linearly within the first 24 h incubation, with rates ranging from 0.003 to 0.01 ml min⁻¹. From those rates we could estimate that the time averaged rate of cilia ventilation should be about 0.16 ml min⁻¹. Since the cilia pumping into the burrow occurs in periods of 24 ± 12 min and at 50-70% of the measured time, considerable amounts of water from deeper sediments may percolate upwards to the sediment surface. This water is rich in reduced compounds and nutrients and may have important associated ecological implications in the ecosystem (e.g. affecting redox conditions, organic matter degradation, benthic recruitment and primary production).     

Depth of artificial Burrowing Owl burrows affects thermal suitability and occupancy

Many organizations have installed artificial burrows to help bolster local Burrowing Owl (Athene cunicularia) populations. However, occupancy probability and reproductive success in artificial burrows varies within and among burrow installations. We evaluated the possibility that depth below ground might explain differences in occupancy probability and reproductive success by affecting the temperature of artificial burrows. We measured burrow temperatures from March to July 2010 in 27 artificial burrows in southern California that were buried 15–76 cm below the surface (measured between the surface and the top of the burrow chamber). Burrow depth was one of several characteristics that affected burrow temperature. Burrow temperature decreased by 0.03°C per cm of soil on top of the burrow. The percentage of time that artificial burrows provided a thermal refuge from above‐ground temperature decreased with burrow depth and ranged between 50% and 58% among burrows. The percentage of time that burrow temperature was optimal for incubating females also decreased with burrow depth and ranged between 27% and 100% among burrows. However, the percentage of time that burrow temperature was optimal for unattended eggs increased with burrow depth and ranged between 11% and 95% among burrows. We found no effect of burrow depth on reproductive success across 21 nesting attempts. However, occupancy probability had a non‐linear relationship with burrow depth. The shallowest burrows (15 cm) had a moderate probability of being occupied (0.46), burrows between 28 and 40 cm had the highest probability of being occupied (>0.80), and burrows >53 cm had the lowest probability of being occupied (<0.43). Burrowing Owls may prefer burrows at moderate depths because these burrows provide a thermal refuge from above‐ground temperatures, and are often cool enough to allow females to leave eggs unattended before the onset of full‐time incubation, but not too cool for incubating females that spend most of their time in the burrow during incubation. Our results suggest that depth is an important consideration when installing artificial burrows for Burrowing Owls. However, additional study is needed to determine the possible effects of burrow depth on reproductive success and on possible tradeoffs between the effects of burrow depth on optimal temperature and other factors, such as minimizing the risk of nest predation.     

Distastefulness as a defense mechanism in Aplysia brasiliana (Mollusca: Gastropoda)

An unusual courtship pattern for fiddler crabs is described from field observations in Panama. This behavior pattern, referred to here as “directing,” differs considerably from the more frequently observed communal courtship system found in close relatives of Uca deichmanni. A male involved in “directing” approaches a female and attempts to carry or maneuver her into his burrow for mating. The female usually struggles to escape from the male. This activity often attracts other males which attempt to “direct” the female if she escapes from the first male. A male is most successful in “directing” a female into his burrow if a) he is larger than the female, b) the female is wandering (a sign of physiological receptivity) prior to the “directing” attempt, and c) several males attempt to “direct” the female at once. The results suggest that females are choosing mates by inciting several males to compete for them. The males which successfully “direct” the struggling females are probably the most fit males.     

Recognition of burrow's olfactory signature in blue petrels, Halobaena caerulea: an efficient discrimination mechanism in the dark

Previous work has shown that blue petrels need olfaction to home. We investigated whether they also recognize an olfactory signature of their own nest. We performed T-maze experiments in which maze arms were connected with the subject bird's burrow and with the burrow of a conspecific neighbour. Of 23 birds, 16 were able to recognize the arm leading to their own burrow. In a second experiment, we positioned the maze in front of the subject's burrow but the maze arms were closed and did not enter the burrow. Consequently, no burrow odours could be sensed by the bird. In this case, 85% of birds (17 of 20) failed to choose, suggesting that petrels were not motivated to choose by positional cues in the absence of odour cues. We explored this idea further by performing a homing experiment whereby homing birds had to relocate an artificially displaced burrow entrance. Blue petrels homed, ignoring the natural burrow entrance and using the new artificial one. The ability to smell their own burrow allows blue petrels to return to the colony at night and to find the correct nest.     

Scatter hoarding by kangaroo rats (Dipodomys merriami) and pilferage from their caches

We observed radio-implanted Merriam's kangaroo rats disposingof 10-g bonanzas of rolled oats in 48 trials in the field. Theprincipal determinant of the initial disposition of discoveredfood was apparently its distance from the day burrow: food foundwithin about 10m was mainly larder hoarded, whereas food encounteredfarther afield was usually dispersed immediately in shallowcaches. Cache sites were newly dug for the purpose and not reused;most caches were nearer the current day burrow than was thefood source, but a few were placed far from both the cacher'sday burrow and its habitual nocturnal range. An experiment withartificial caches indicated that security from discovery increaseswith spacing and with proximity to perennial shrubs. Nine kangaroorats cached dyed food, and fecal dye traces revealed extensivepilferage from five of them, by both conspecifics and otherrodent species. Limited evidence indicates that food encounterednearer home and initially larder hoarded was more secure frompilferage than food initially scattered, and yet kangaroo ratswere observed to scatter caches soon after initial larder hoarding.A kangaroo rat whose dyed stores escaped pilferage fed fromthem at intervals for at least 12 days. Even cachers who incurredpilferage made as much, or more, use of their caches as anythief, suggesting that scattering caches may be a defense againstcatastrophic losses.     

Division of Labour and Extended Parenting in a Desert Tenebrionid Beetle

Parastizopus armaticeps (Peringuey) is a nocturnal detritivorous desert tenebrionid beetle which can only reproduce after heavy rainfall. Both parents care for the young and excavate a breeding burrow together. They play similar roles in burrow excavation and protection until the eggs hatch, after which male and female roles diverge markedly Females collect detritus on the surface at night, depositing this at the burrow entrance, which they also clear of sand in the early evening. Males remain inside the burrow and dig to deepen and extend it, maintaining its moisture level as the sand desiccates They also pull the food dropped by the female down to the burrow base to form a food store for the offspring. Both sexes guard the burrow and attack same-sex intruders, especially in the early phases of reproduction. The parents remain inside with the pupal cocoons until the teneral adults eclose. Males dig continually to maintain burrow humidity while females, which rarely forage during the pupal Period, recommence foraging for a further 16-18d after pupal eclosure. They provide food for the newly eclosed beetles until the exoskeletons have hardened and melanization is almost complete, then the family disperses. Offspring over 10 d old pull down forage at the burrow entrance and later forage for food themselves and carry it to the burrow. This is the highest level of social and reproductive complexity described for beetles and the behavioural and ecological factors contributing to its evolution are discussed in the context of parental care known for other subsocial arthropods, especially coleopterans.     

Earthen Structures Built by Ilyoplax dentimerosa (Crustacea,Brachyura, Ocypodidae)

Under observation, the small ocypodid crab Ilyoplax dentimerosa was found to commonly build three types of earthen structures: a barricade near its neighbour's burrow, a fence at an intermediate position between the burrows of the builder and its neighbour, and a minishelter near the builder's burrow. The sex ratio of barricade builders was found to be close to 1:1, whereas most of the fence builders were found to be female. Crabs against which barricades and fences were built, were usually smaller than the builders. Both barricade builders and fence builders had, in most cases, minishelters at the side, facing the barricade or the fence. Removal and rebuilding experiments for barricades and fences demonstrated that both structures functioned to deter the approach of the builder's neighbour to the builder's activity site. Fences also had the effect of restraining the movement of the builder toward the fence site.     

Homing in fiddler crabs (Uca lactea annulipes and Uca vomeris : Ocypodidae)

Fiddler crabs emerge from burrows on intertidal sand- and mudflats to feed during low tide. In the species studied here (Uca lactea annulipes, Uca vomeris) a crab normally wanders no more than about 1 m away from its burrow and, when frightened, dashes back along a straight line to take cover. Feeding crabs tend to move sideways, without changing orientation, along paths radiating from the burrow. When they move along circumferential paths they adjust their orientation so that one side continues to point towards the burrow. The crabs do not need to see the burrow in order to stay aligned with the home vector, and they are not misled by a dummy hole close to their own burrow unless they have come to within about 10 cm of it. The home runs of crabs end within a few centimeters of a burrow that is covered with a sheet of sandpaper and then give way to search runs, centred upon a position slightly short of the burrow location. Feeding crabs can be displaced on sandpapers and their subsequent home runs end at a position where the burrow would be, had there been no displacement. Landmarks close to the burrow do not influence the home runs of displaced crabs. Crabs that are rotated on a sheet of sandpaper, counter-turn to keep their original orientation constant. Fiddler crabs thus employ path integration with external compass information and close range visual guidance for homing. Accepted: 11 May 1998