The underground life of the oriental mole cricket: an analysis of burrow morphology

The underground life of the oriental mole cricket Gryllotalpa orientalis has been investigated by studying the structure of its burrows under different environmental situations and in different seasons. The different uses of different burrow types and their advantages and disadvantages have been examined. The total length, number of tunnels and combination of burrow types varied from a simple tunnel to a more complex one with branches at various angles to the surface, burrow types being divided roughly into shallow horizontal or deep vertical ones. In horizontal burrows, the branching structure was well developed in various directions. It is notable that the vertical burrows of G. orientalis were occupied by only one individual. Both vertical and horizontal burrows were used for foraging: vertical burrows for plants with subterranean stems and horizontal burrows for creeping plants. Vertical burrows were also used for hiding from predators, resting, moulting and overwintering, whereas horizontal burrows were used for escaping from predators and as mating routes. Egg chambers were constructed beside horizontal burrows, and calling burrows were constructed as part of horizontal burrows. Based on their current requirements, mole crickets continuously modify their burrow structures or change burrowing sites.     

Burrow architecture and digging activity in the Cape dune mole rat

While females are traditionally thought to invest more time and energy into parental care than males, males often invest more resources into searching and displaying for mates, obtaining mates and in male–male conflict. Solitary subterranean mammals perform these activities in a particularly challenging niche, necessitating energetically expensive burrowing to both search for mates and forage for food. This restriction presumably affects males more than females as the former are thought to dig longer tunnels that cover greater distances to search for females. We excavated burrow systems of male and female Cape dune mole rats Bathyergus suillus the, largest truly subterranean mammal, to investigate whether male burrows differ from those of females in ways that reflect mate searching by males. We consider burrow architecture (length, internal dimensions, fractal dimension of tunnel systems, number of nesting chambers and mole mounds on the surface) in relation to mating strategy. Males excavated significantly longer burrow systems with higher fractal dimensions and larger burrow areas than females. Male burrow systems were also significantly farther from one another than females were from other females' burrow systems. However, no sex differences were evident in tunnel cross-sectional area, mass of soil excavated per mound, number of mounds produced per unit burrow length or mass of soil excavated per burrow system. Hence, while males may use their habitat differently from females, they do not appear to differ in the dimensions of the tunnels they create. Thus, exploration and use of the habitat differs between the sexes, which may be a consequence of sex differences in mating behaviour and greater demands for food.     

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.     

Effects of burrow condition and seed handling time on hoarding strategies of Edward's long-tailed rat (Leopoldamys edwardsi)

Many hoarding rodents use burrows not only for dwelling and protection from natural enemies, but also for food storage. However, little is known how burrows used by scatter-hoarding animals influence their foraging behaviors. In addition, handling time for a given food item has a fundamental impact on hoarding strategies of these hoarding animals: food items with longer handling time are more likely to be hoarded due to increasing predation risk because the animals spend more time outside their burrows if they consumed such food. By providing with two types of artificial burrows (aboveground vs. underground) and two types of food items (i.e. seeds) with contrasting handling times, we investigated how burrow condition and handling time co-influence hoarding strategies of a key scatter-hoarding rodent, Edward's long-tailed rat (Leopoldamys edwardsi) in large enclosures in southwest China. We found that only a few animals larder-hoarded fewer seeds when only aboveground burrows were available, while over 80% of the animals preferred to use the underground burrows and hoard significantly more seeds in the burrows when both aboveground and underground burrows were provided simultaneously. We also found that seed handling time significantly affected hoarding strategies of the animals: they consumed and/or scatter-hoarded more Camellia oleifera seeds with shorter handling time outside the burrow, but consumed and larder-hoarded more Lithocarpus harlandii seeds with longer handling time in underground burrows. Our study indicates that both burrow types and seed handling time have important impacts on hoarding strategies of scatter-hoarding animals.     

Orientation of crustacean burrows in the Bay of Panzano (Gulf of Trieste, Northern Adriatic Sea)

Three burrow systems produced by Jaxea nocturna and one cluster of burrows produced by Upogebia pusilla where investigated in the Bay of Panzano, Northern Adriatic Sea, to determine preferred orientations. The distributions of dip directions differ between both producers. Steep shafts into the consolidated mud followed by large, shallowly inclined tunnels oriented in a manner similar to a spiral are characteristic for Jaxea nocturna burrows. In contrast, the Y-shaped burrows of Upogebia pusilla have entrance shafts that are less steep which are connected by a near-horizontal section, where a blind ending tunnel branches. Orientations of the dominating, shallowly inclined burrow parts are not randomly distributed in either Jaxea or in Upogebia burrows. Three preferred axial orientations with almost equal proportions in each direction are characteristic for the Jaxea burrow from the centre of the bay. This system transforms to a squared structure near the coast, where one direction parallels the shoreline and the other is oriented at right angles to the coast. The latter orientation demonstrates significant constancy in all investigated burrows. Upogebia burrow clusters coincide with the three preferred orientations of the Jaxea burrow from the bay centre, but the proportions of the directions are unequal. Burrow segments connecting the steep entrance shafts in Upogebia are oriented almost parallel to the shoreline, whereas at right angles to the coast the flat blind-ending tunnels incline towards the open sea. The study shows a strong coincidence between empirical and theoretical distributions of the dominating orientations in both species.     

Orientation of crustacean burrows in the Bay of Panzano (Gulf of Trieste, Northern Adriatic Sea)

Three burrow systems produced by Jaxea nocturna and one cluster of burrows produced by Upogebia pusilla where investigated in the Bay of Panzano, Northern Adriatic Sea, to determine preferred orientations. The distributions of dip directions differ between both producers. Steep shafts into the consolidated mud followed by large, shallowly inclined tunnels oriented in a manner similar to a spiral are characteristic for Jaxea nocturna burrows. In contrast, the Y-shaped burrows of Upogebia pusilla have entrance shafts that are less steep which are connected by a near-horizontal section, where a blind ending tunnel branches. Orientations of the dominating, shallowly inclined burrow parts are not randomly distributed in either Jaxea or in Upogebia burrows. Three preferred axial orientations with almost equal proportions in each direction are characteristic for the Jaxea burrow from the centre of the bay. This system transforms to a squared structure near the coast, where one direction parallels the shoreline and the other is oriented at right angles to the coast. The latter orientation demonstrates significant constancy in all investigated burrows. Upogebia burrow clusters coincide with the three preferred orientations of the Jaxea burrow from the bay centre, but the proportions of the directions are unequal. Burrow segments connecting the steep entrance shafts in Upogebia are oriented almost parallel to the shoreline, whereas at right angles to the coast the flat blind-ending tunnels incline towards the open sea. The study shows a strong coincidence between empirical and theoretical distributions of the dominating orientations in both species.     

Laboratory observations on the burrowing of the snake blenny, Lumpenus lampretaeformis (Walbaum), in soft sediment

The Lumpenus lampretaeformis used in this study were caught on soft sediments in Irvine Bay and Ardmucknish Bay (West Coast of Scotland). In aquaria the fish is capable of constructing and maintaining its own burrow in soft sediments. The burrow has a'Y'configuration which may be made complex by the construction of additional tunnels as the original system begins to decay. The cross sections of these burrows are distinctive in that they have an oval shape in the main tunnel and a rounded cross section in the subsidiary (side) tunnel. Burrowing can be relatively rapid, a burrow with three entrances being constructed within 12 h.     

Evolution of African mole-rat sociality: burrow architecture, rainfall and foraging in colonies of the cooperatively breeding Fukomys mechowii

African mole rats (Bathyergidae) offer an excellent system with which to test theories relating to the evolution and maintenance of sociality in mammals. The aridity food distribution hypothesis (AFDH) suggests that, within the bathyergids, sociality has evolved in response to patterns of rainfall, its effects on food distribution, and the subsequent costs and risks of foraging and dispersal. Here, in the first detailed study of burrow architecture in a social mole-rat species, with data from 32 burrows, we show that in the giant mole-rat Fukomys mechowii burrow fractal dimension increases with colony size and is higher during the rainy season than during the dry season. The mass of food in the burrow increases with fractal dimension and is higher during the rainy season than during the dry season. These results link for the first time colony size, burrow architecture, rainfall and foraging success and provide support for two assumptions of the AFDH, namely that (1) in arid conditions burrowing may be severely constrained by the high costs of digging; and (2) the potential risks of failing to locate food may be mitigated by increases in colony size.     

Burrow fractal dimension and foraging success in subterranean rodents: a simulation

For animals that forage underground, the success with whichfood items are located may be closely related to burrow architecture.Fractal dimension, which describes how a burrow explores thesurrounding area in a way that is independent of burrow length,is an obvious choice for a single metric describing burrow shape.Although it is often assumed that burrows of high fractal dimensionwill be associated with greater foraging success, this has notpreviously been demonstrated. In this study, we use computersimulations to study the success with which burrows of differentfractal dimensions locate randomly distributed food items. Inaddition, we examine the effect of different patterns of fooddistribution (in particular the patchiness with which food itemsare distributed) and consider how using different criteria forlocating food items affects the relationship between fractaldimension and foraging success. We conclude that, under a widerange of plausible assumptions about the ways in which subterraneanrodents forage, burrows of high fractal dimension are more successfulat locating food items than burrows of lower fractal dimension.     

Burrow use by Tibetan Ground Tits Pseudopodoces humilis: coping with life at high altitudes

The Tibetan Ground Tit Pseudopodoces humilis is a high-altitude passerine endemic to the Tibetan Plateau. A 4-year study in alpine meadows in Northern Tibet at 4300 m asl demonstrated that rather than using Pika Ochotona spp. holes as previously reported, the birds excavated one nest burrow themselves in spring and another after breeding, which they used for roosting in winter. Both the nesting and winter-roosting burrows comprised a straight tunnel leading to an ellipsoid chamber. There were no significant differences in placement or structural characteristics between the two types of burrows, except that winter-roosting burrows had a significantly smaller entrance diameter. Most burrows were 100–160 cm long and their chambers 20–40 cm deep. Tibetan Ground Tits tended to maximize the thermal benefits of their burrows by adjusting their spatial and structural characteristics in response to local solar radiation and wind regimes. Burrows tended to be oriented towards the sun and away from prevailing winds, presumably to maintain burrow temperature. Longer tunnels could function to save heat from solar radiation or reduce wind-disturbance, while shorter tunnels allow chambers to be warmed sooner in situations where wind potentially reduced soil temperatures. The thermal benefits to the birds of burrow architecture are likely to play a crucial role throughout the year in these extreme alpine environments.     

The burrow ofDasypus hybridus (Cingulata: Dasypodidae)

Dasypus hybridus (Desmarest, 1804) inhabits open fields in southern South America. Burrows ofD. hybridus were cylindrical with a conical end. They had an entrance and a single tunnel without branches. Three locations of the burrows in the terrain were detected: in banks of dried waterways, near rocks, and in open field. The orientation of 29 burrow mouths was not random and, although they did not point to any cardinal point, in particular there was a strong tendency to avoid the south quadrant. In a sample area of 90 × 140 m, burrow mouths were arranged in a random spatial pattern with a density of 25.4 burrows per ha. Great variation in burrow length was found (118.8 ± 105.69 cm, CV = 89.0%). Tunnels may be used as refuges and/or for thermoregulation. The orientation of burrow mouths can also be related to thermoregulation as tunnels are covered from dominant winds, and, for many hours every day, the temperature at the mouths can be influenced by insolation. The possible existence of shelter-burrows and resting-burrows is discussed.     

Burrow morphology and utilization of the goby (Parapocryptes serperaster) in the Mekong Delta,Vietnam

Some fish species living in mudflats construct burrows for dwelling and hiding. The goby Parapocryptes serperaster is a burrowing fish in mudflats of many estuaries in South East Asia. This study was carried out in the Mekong Delta, Vietnam, to examine burrow morphology and usage by this species. Morphology of the burrows constructed by P. serperaster was investigated by resin castings in situ to obtain the physical structure and configuration of each burrow. Fish from the burrows were caught and measured before burrow casts were made. Fish burrows comprised several openings, a few branching tunnels and multi-bulbous chambers. The surface openings were circular, and the shapes of branching tunnels were nearly round. The burrows had interconnected tunnels and various short cul-de-sac side branches. The burrow structure differed between fish sizes, but burrow dimensions were positively correlated with fish size, indicating that larger fish can make larger and more sophisticated burrow. The burrow structure and dimensions were not different between the dry and wet seasons. Laboratory observations showed that P. serperaster used body movements to dig burrows in the sediment. Burrows could provide a low-tide retreat and protection from predators, but were not used for spawning and feeding for this goby species. This study indicates that the burrowing activity of gobies is an important adaptation for living in shallow and muddy habitats.     

Pattern of Dispersion of Young Wild Rabbits,Oryctolagus cuniculus L., in Relation to Burrows

The development of dispersion in relation to burrows of young rabbits, Oryctolagus cuniculus L., was studied in a sand dune habitat between May and September 1984–1985. Generally, young rabbits did not show a close association with their original burrow. From the first week of life on the surface they used different burrows as well as the original one. No significant age-related changes in the mean distance from different kinds of burrows were observed. The mean distance from the nearest burrow remained always under 3 m, but this distance may have been due largely to the high density of burrows. The apparent freedom of movements of young rabbits around different burrows may be related to the social system of the adults in a sand dune habitat.     

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

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

Advanced sub social behaviour in the scorpionHeterometrus fulvipes Brunner (Arachnida)

Scorpions arc generally non-social, solitary animals that interact with conspecifics at birth, courtship or predation only. The present study reports the presence of advanced sub social behaviour inHeterometrus fulvipes Brunner and evaluates the importance of its burrowing as a cause for such social behaviour.Heterometrus fulvipes constructed deep angular burrows at the base of plants. Burrows provided (i) protection against predation, (ii) increased availability of food and (iii) ideal microclimate for year round activity of the scorpions. No cannibalism was observed in laboratory maintained colonies. The risk of predation and the difficult by immatures to dig tunnels during dry soil conditions may have forced the mother and offspring to live together in the burrow for longer durations. The cohabitation of relative offsprings transforms the burrow into a nest. The members of a colony exhibits division of labour for nest expansion and in foraging. The mother communicates with the immatures through “Buzz” sound and may provide premasticated food. There is food sharing also among colony members. All these behaviours indicate the presence of advanced sub social behaviour inHeterometrus fulvipes.     

Responses of the endangered pygmy bluetongue lizard to conspecific scats

Many animals use chemical signals for communication between conspecifics and for territory marking. The pygmy bluetongue lizard is normally solitary, focussing activity around the entrance of its burrow, from where it ambushes prey, and rarely contacts other individuals. In this paper we examined whether lizards in laboratory experiments alter their behaviour in the presence of scats from conspecifics. In the first experiment, when lizards were offered a choice of two vacant burrows with or without a scat close to the entrance, they tongue flicked more often at the burrow entrance when the scat was present, and more often chose to occupy the burrow with the scat. An interpretation is that lizards use scat signals to recognise burrows that may be suitable because they have previously been occupied by a conspecific, but that they approach those burrows cautiously in case a resident is still present and likely to resist a takeover. Scats from male lizards were inspected (by both sexes) for longer than scats of female lizards. In the second experiment, when resident lizards were presented with scats outside of their burrows, they inspected and tongue flicked at those scats more often if the scat came from a male than a female lizard, but there was no definitive evidence from our experiments that lizards differentiated in their response to scats from lizards that were found close to or far from the test lizard. The results were consistent with a communication system in which lizards use scats to advertise their presence, independent of any direct contact.     

Burrow morphology and dynamics of mudshrimp in Asian soft shores

Mudshrimps are important soft shore bioturbators but research on the ecology of tropical species has received less attention when compared with their temperate counterparts. The mudshrimp Austinogebia edulis is common on Asian soft shores and lives in burrows for its entire adult life. Epoxy resin casting of A. edulis burrows showed that they were approximately Y-shaped, with an upper U-part and the lower central shaft part. The burrows had two openings extending to the surface; the mean distance between the two openings was 11.0 cm in Hong Kong and 26.4 cm in Taiwan. Openings of the burrows had small chimneys. The tunnels of the burrows were circular, narrow and with a smooth surface (tunnel diameter corresponded to shrimp carapace width). Each burrow was inhabited by a single shrimp and burrows were inter-connected during the mating and reproductive season. Each burrow had four to 12 spherical chambers, which were free of detritus. The chambers were thought to be used for suspension feeding, current generation and as turning points. The depth of burrows was up to 1.1 m. Multivariate analysis on various burrow parameters showed that burrows collected on a mud flat in Taiwan were deeper, had a wider distance between the openings and a larger volume than burrows collected from a sandy shore in Hong Kong, suggesting that burrow architecture is variable between shore types. Burrow architecture, however, did not vary between tidal levels, seasons and shrimp density on the shores in Hong Kong, indicating that the burrows were quite stable within the substratum and were not affected by environmental and biological factors.     

Burrow plasticity in the deep-sea isopod Bathynomus doederleini (Crustacea: Isopoda: Cirolanidae)

We investigated whether the deep-sea isopod Bathynomus doederleini has the capacity to change burrow length in response to changes in environmental conditions. We observed burrowing behavior in individuals that were placed on substrates with either simple (ST) or complex (CT) surface topographies. Individuals in the ST group (N = 10) constructed seven burrows. The mean ratio of the burrow length to body length was 1.8. The individuals in the CT group (N = 10) constructed eight burrows with a mean ratio of burrow length to body length of 2.5. Thus the burrows were significantly longer in the CT group. In addition, the isopods in the CT group often incorporated a chamber in the mid-section of the burrow. Our results may be used to infer the determinants of burrow morphology and speculate about the lifestyle of this species in the deep sea.     

Description of the nursery burrow of the Mexican cottontail rabbit Sylvilagus cunicularius under seminatural conditions

The rabbit Sylvilagus cunicularius is endemic to Mexico and is one of the largest, most widespread, but little studied of the country’s 10 rabbit species. As part of a project investigating its reproductive biology, we describe here for the first time the nursery burrow, vital for the survival of the altricial young. During the breeding seasons of 2008–2012, we collected data from 25 nursery burrows and 22 nests constructed by eight females (three wild caught and five captive bred) kept in enclosures within their natural habitat in the Malinche National Park in the central Mexican state of Tlaxcala. Although not a burrow-living species, several days before parturition, females dug a nursery burrow in which they constructed a nest. These burrows were short, shallow tunnels with a median length of 23 cm, typically located beneath grass tussocks or shrubs, and ending in a nest chamber a median 17 cm beneath the surface. Nests consisted of dry grass, fragments of woody plants, pine needles, and alfalfa hay and oat straw provided as supplementary food, fur pulled from the mother’s body, and her fecal pellets. Females nursed their young at the burrow entrance, and until the young were approximately 12 days old, they closed the entrance after each visit in such a way as to make it very difficult for humans to locate. Surprisingly, these nursery burrows were more similar to those of the European rabbit Oryctolagus cuniculus and the pygmy rabbit Brachylagus idahoensis than to other Sylvilagus species studied to date. As almost 50 % of burrows did not result in emergent young, present work is directed to determining what characterizes successful burrows.     

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.