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.     

Home-Range Dynamics in a Solitary Subterranean Rodent

Despite an important role of subterranean rodents as ecosystem engineers, their belowground mobility is poorly documented. It is supposed that their underground burrow systems, once established, are relatively stable because of high-energy costs of digging. We chose the silvery mole-rat, Heliophobius argenteocinereus (Bathyergidae, Rodentia) from mesic Afrotropics as a representative of solitary subterranean rodents to investigate how, and how fast these rodents process their established burrow systems. We combined radio-tracking of individual animals with subsequent mapping of their burrow systems, and we developed a new method for assessing the rate of burrowing. Mole-rats continuously rebuilt their burrow systems; they excavated approx. 0.7 m of new tunnels per day and backfilled on average 64% of all tunnels. On average, every 32 d they established a new nest. They often completely backfilled newly excavated peripheral burrows, while other parts of their burrow systems were more permanent. Their home-ranges were dynamic and continuously shifted in space. Burrow system processing continued even in the advanced dry season, when soil is difficult to work.     

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.     

What determines the way of deposition of excavated soil

Subterranean rodents continuously extend their burrow systems primarily in search of food, which has an important impact on the ecosystem in which they live. Excavated soil may be pushed either into aboveground mounds or into tunnels underground. Factors affecting the amount of burrowing and the preference of aboveground or underground soil deposition are, nevertheless, little known. We investigated the influence of food supply, soil hardness, and the animal’s body mass on the mode of soil deposition in ten burrow systems of free ranging silvery mole-rats Heliophobius argenteocinereus Peters, 1846. In each burrow system, we estimated the volume of backfilled tunnels and the volume of soil deposited aboveground. The highest amount of variation in these parameters was explained by the interaction of food supply and soil hardness. The ratio of the volume of backfilled tunnels to the volume of mounds was not significantly dependent on any of the explanatory variables. The proportion of backfilled tunnels decreased with the increasing volume of the complete burrow system. We propose that both low food supply and soft soil lead to an increased amount of burrowing, which results in a larger volume of soil deposited both above ground and under ground over a given period of time.     

Area-restricted search by the plains pocket gopher (Geomys bursarius) in tallgrass prairie habitat

Because pocket gophers have the high energetic cost of excavatingburrows and an inability to detect distant food items throughthe soil, I hypothesized that individuals within establishedburrow systems would use area-restricted search as a foragingstrategy. To examine this hypothesis I compared gopher foragingeffort over a 10-month period between areas in which overallplant densities were experimentally varied. Gophers expendedapproximately 50% of their foraging effort in areas with thehighest plant density, even though these made up only 33% ofthe available area in experimental plots. In large, griddedareas sampled for an entire season as well as in small areasin which gophers foraged for less than 1 week, gopher foragingeffort was related to the density of a single leguminous plantspecies, Psoralea argophylla. In small plots where this plantspecies was at high density, gophers created more tunnel branches,thereby intensifying their search effort. Thus, area-restrictedsearch appears to increase the rate of encounter with the patchilydistributed Psoralea plants.     

Burrow patterns of the mole rat Spalax ehrenbergi in two soil types (terra-rossa and rendzina) in Mount Carmel, Israel

Complete burrow systems of the mole rat Spalax ehrenbergi (Rodentia; Spalacidae), from two sites on Mount Carmel in Israel, with different soil types (terra-rossa and rendzina), were excavated and described here for the first time.
A comparison was made of burrow structure patterns in the two soils and of the sexes, with special attention to the features of the feeding tunnels.
The pattern in the rendzina revealed longer burrows with a longer main tunnel and fewer branches per metre of the main tunnel, while the pattern in the terra-rossa revealed shorter burrows with a shorter main tunnel and a relatively higher number of branches. These differences can primarily be related to the different levels of food availability, which is higher in the terra-rossa. It is suggested that each of the patterns reflects the mole rat's ability to optimize foraging efficiency in accordance with its given food availability.
The average total length of the males' burrows was much greater than those of the females' burrows in the rendzina soil. It appears that food requirements determine different burrow features of the sexes rather than reproduction requirements.
Other tunnel features (e.g. structural complexity, depth and width, segment length and spatial arrangement) and the factors which may affect them, as well as burrow structure of young mole rats and evidence of the underground dispersion of young mole rats, were presented and discussed.
Many similarities were found in a comparison of S. ehrenbergi burrow features with those of other solitary subterranean rodents.     

Postnatal development of subterranean habits in tuco-tucos <Emphasis Type="Italic">Ctenomys talarum</Emphasis> (Rodentia,Caviomorpha, Ctenomyidae)

Postnatal development of subterranean habits was investigated in Los Talas’ tuco-tuco Ctenomys talarum, a subterranean caviomorph rodent endemic from South America. Since in this species, some key morpho-functional traits related to scratch-digging behaviour—a form of underground progression—are already present during early ontogeny and develop progressively, we predicted that this behaviour expresses early during postanatal development and its performance enhances gradually from pups to adults. The process of acquisition of different behaviours associated to the construction of a burrow system was recorded in 11 individuals, each one coming from different litters, inside a terrarium filled with natural soil. We found that scratch-digging and burrowing behaviours expressed early during postnatal development, particularly, during lactancy. The digging of a “true burrow” clearly preceded the dispersal age, with a high inter-individual variability, from 18 (lactancy) to 47 (post-weaning) postnatal days. Pups could lose the soil using their foreclaws and remove the accumulated substrate using their hindfeet as adults do. During lactancy individuals could construct a simple burrow to shelter, and first burrow construction occurred in the absence of either a burrowing demonstrator or an early subterranean environment (a natal burrow). However, certain features of the complex burrow system that characterize this species, such as lateral branches and nest chamber, just appeared after weaning. The time elapsed until animals started to dig and the time dedicated to underground activities varied with age, decreasing and increasing, respectively. In sum, our results show that—in C. talarum—immature digging behaviour gets expressed early during ontogeny, and develops progressively. The role of the early ability to build its own burrow and its possible function influencing the development of musculoskeletal traits and on efficiency for such conduct is discussed.     

祁连山东段高原鼢鼠采食洞道气体研究

研究高原鼢鼠（Myospalax baileyi）洞道气体环境对了解其低氧适应机制具有重要作用。本试验在祁连山东段高寒草甸利用土壤原位气体测定仪,对高原鼢鼠采食活动洞道、非活动洞道、地表空气及无洞道土壤内的温度、氧气、二氧化碳、甲烷含量进行了连续12个月的监测。通过One-Way ANOVA检验、重复测量方差分析以及Pearson相关性分析发现,（1）高原鼢鼠活动洞道氧气含量,除10月、11月与非活动洞道无显著性差异外（P>0.05）,均显著小于非活动洞道、地表空气和无洞道土壤（P<0.05）;活动洞道二氧化碳含量,除7月、8月份与非活动洞道无显著性差异外（P>0.05）,均显著大于非活动洞道、地表空气和无洞道土壤（P<0.05）;活动洞道内甲烷含量各月均显著高于地表空气甲烷含量（P<0.05）,与非活动洞道、无洞道土壤的月季差异各异。（2）高原鼢鼠活动洞道内氧气含量的最小值和二氧化碳的最大值均出现在5月和9月,其它处理下氧气最小值和二氧化碳最大值均出现在6-8月;4个处理下甲烷含量最小值在1月与12月出现,最大值出现在5月和9月。（3）月份、处理以及月份和处理间的交互作用均对氧气、二氧化碳、甲烷含量有显著影响（P<0.05）。可见,高原鼢鼠生存在低氧、高二氧化碳和较高甲烷含量的环境中,且洞道内部的气体环境会受季节和高原鼢鼠活动的影响。     

Influence of soil compaction on tunnel network construction by the eastern subterranean termite (Isoptera: Rhinotermitidae)

Eastern subterranean termites, Reticulitermes flavipes (Kollar), workers were introduced into arenas containing low, moderate, and high compaction builder's sand (1.05 g/cm3, 1.18 g/cm3 or 1.35 g/cm3 bulk densities, respectively), and they immediately began tunneling. Termites built the tunnel network significantly fastest in soil of low compaction compared with moderately or highly compacted soil. In soil of low compaction, 221.67 +/- 4.73 cm of total tunnel distance was constructed in 1 d compared with only 96 cm of tunneling in highly compacted soil. At 14 d, total tunnel distance averaged 216.83 +/- 4.56 cm in soil of low compaction compared with 169.70 +/- 4.10 and 181.18 +/- 6.13 cm in moderately and highly compacted soil, respectively. Decreases in total tunnel distance between 1 and 14 d were caused by backfilling of seldom-used tunnels. Termites did the majority of tunneling during the first day of introduction into arenas. In soil of low and moderate compaction, termites essentially constructed the entire tunnel network within the first day, only modifying it by backfilling or maintaining tunnels. In highly compacted soil, 53% of the final tunnel network was constructed during the first day, 87% was constructed by the third day, and 97% was constructed by the seventh day. Soil compaction did not affect the number of primary tunnels or the number and diameter of secondary tunnels. The angle between the secondary tunnel and primary tunnel also was not significantly affected by soil compaction. However, the number of secondary tunnels in soil of low compaction (5.89 +/- 0.51) was significantly greater than in moderately (2.74 +/- 0.36) and highly (3.58 +/- 0.59) compacted soils.     

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.     

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.     

Seasonal changes in body composition of Ctenomys talarum (Rodentia: Octodontidae): an herbivore subterranean rodent

Ctenomys talarum is a subterranean herbivorous rodent whose burrow systems exhibit particular characteristics, distinct from other subterranean environments. We studied seasonal variation in body composition of C. talarum in relation to energetic requirements. Body lipid content seasonally changed in C. talarum, related to reproductive cycle and thermorregulatory mechanisms. A decrease in protein body content was found only in spring. Ash content of females was lowest when most of them are in post partum estro. Observed variations in water body content could be associated with plant water content and/or metabolic regulation. Our results show the occurrence of seasonal variations in body composition in C. talarum, which could be related to the high cost of reproduction and the subterranean life style of this species.     

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.     

Tunnel formation by Reticulitermes flavipes and Coptotermes formosanus (Isoptera: Rhinotermitidae) in response to wood in sand.

The tunneling responses of two subterranean termite species, Coptotermes formosanus Shiraki and Reticulitermes flavipes (Kollar), to the presence of sound wood in laboratory arenas were studied. Branching pattern and the speed of tunnel construction between R. flavipes and C. formosanus also were compared. Patlak's residence index (rho) was generated using the length, width, speed of construction, and area of the primary tunnels built by termites. In the same allotted time, C. formosanus built wider and shorter primary tunnels, whereas R. flavipes built thinner and longer primary tunnels. The presence of wood did not affect termite tunnel formation. This lack of variation in tunnel formation parameters was evidenced by the inability of the termites to locate wood sources over distance, even as short as 2.5 mm, and by the similar tunneling behaviors in areas of the arena with or without wood. Patlak's model predicted the densities of tunnels with an error between 9 and 28%. in experiments with R. flavipes exposed to a range of 0-8,000 g of wood, and between 61 and 87% in experiments with C. formosanus. These results indicated that the residence index can provide a qualitative measure of the effect of habitat heterogeneity on the individual termite tunnels. The tunneling constructions strategy of these subterranean termites is discussed.     

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.     

The influence of foraging mode and arid adaptation on the basal metabolic rates of burrowing mammals

Two competing but nonexclusive hypotheses to explain the reduced basal metabolic rate (BMR) of mammals that live and forage underground (fossorial species) are examined by comparing this group with burrowing mammals that forage on the surface (semifossorial species). These hypotheses suggest that the low BMR of fossorial species either compensates for the enormous energetic demands of subterranean foraging (the cost-of-burrowing hypothesis) or prevents overheating in closed burrow systems (the thermal-stress hypothesis). Because phylogentically informed allometric analysis showed that arid burrowing mammals have a significantly lower BMR than mesic ones, fossorial and semifossorial species were compared within these groups. The BMRs of mesic fossorial and semifossorial mammals could not be reliably distinguished, nor could the BMRs of large (>77 g) arid fossorial and semifossorial mammals. This finding favours the thermal-stress hypothesis, because the groups appear to have similar BMRs despite differences in foraging costs. However, in support of the cost-of-burrowing hypothesis, small (<77 g) arid fossorial mammals were found to have a significantly lower BMR than semifossorial mammals of the similar size. Given the high mass-specific metabolic rates of small animals, they are expected to be under severe energy and water stress in arid environments. Under such conditions, the greatly reduced BMR of small fossorial species may compensate for the enormous energetic demands of subterranean foraging.     

西南岩溶槽谷区隧道建设的水文生态环境效应研究进展

西南岩溶槽谷区隧道分布密集,隧道突水引发了一系列的生态环境问题。梳理了隧道建设产生的生态环境效应现状,包括:改变水资源分布格局及水文过程,诱发地质灾害,降低土地质量和引起土壤污染,影响植被生长与分布等。在隧道影响水文和水资源方面已有丰富的研究成果,而在对土壤和植被的影响研究方面缺乏统一认识。指出了未来研究的总体趋势和方向:隧道影响水资源分布格局与水文过程的水文地质模式,降水、地表水、土壤水、地下水及隧道水的转换过程与地下水流场演化机制,岩溶隧道区植被生理过程与多样性变化等。     

Effects of concentration,distance, and application methods of Altriset (Chlorantraniliprole) on eastern subterranean termite (Isoptera: Rhinotermitidae)

The effects of various concentrations, distance, and application methods of Altriset (Chlorantraniliprole) were investigated against one of the most destructive termites, the eastern subterranean termite, Reticulitermes flavipes Kollar. Three laboratory experiments were conducted. First, we examined the concentration effect of treating the soil contiguously to established foraging tunnels at a fixed 1 m distance. The results demonstrated 100% termite control in 19 d posttreatment at 100 and 50 μg/g and 27% termite mortality at 25 μg/g. Second, we tested the distance effect of the soil treatment (2 and 4 m) on the efficacy of Altriset to the satellite termite populations at a fixed 50 μg/g concentration. This resulted in 100% termite control in 22 d posttreatment at both 2 and 4 m. Third, we examined the effect of differing application methods using 12.5 and 25 μg/g prior to the establishment of foraging tunnels at a fixed 1m distance. This illustrated 100% termite control in 9 d posttreatment at 25 μg/g and 12 d posttreatment at 12.5 μg/g. The third experiment demonstrated soil treatments that were applied prior to termite tunnel establishment had greater efficacy than applications made post tunnel construction. Our results provide a comprehensive understanding about the efficacy of Altriset treatments on eastern subterranean termites.     

The burrow system of the African ice rat Otomys sloggetti robertsi

We studied the architecture of the burrow system of the African ice rat Otomys sloggetti robertsi, a non–hibernating, diurnal murid rodent endemic to the sub–alpine and alpine regions of the southern African Drakensberg and Maluti mountains. In our study site we found ice rat burrows in two substrates (organic and mineral soils). The structure of the burrow system was similar in both soil types, comprising several interlinking tunnels, numerous burrow entrances and 1–2 nest chambers. However, the surface area of the burrow systems in organic soils was larger, the tunnels were deeper, and some of the systems contained two levels, all of which was contrary to our assumption that digging would be more difficult in the compact organic soils. Ice rats occur in colonies of up to 17 individuals, and the collected efforts of several individuals are required for constructing complex burrow systems. The burrow structure is similar to those of two arid–adapted relatives, Parotomys brantsii and Parotomys littledalei, suggesting that the burrow architecture among these three taxa may reflect the similar functions of burrows in extreme environments. For ice rats, burrows could provide a suitable microhabitat in which to escape adverse environmental conditions, particularly during winter. Moreover, ice rat burrows contained far fewer nest chambers than those of both Parotomys species, indicating that members in a colony share nest chambers, thereby facilitating huddling. Finally, the extensive interlinking tunnels may provide underground routes to aboveground feeding sites, thereby reducing exposure to adverse conditions.     

Use of vegetation chemical signals for digging orientation in the subterranean rodent Ctenomys talarum (Rodentia: Ctenomyidae)

Sensory modalities involved in the localization and selection of food represent critical information for understanding the foraging behavior of subterranean rodents. The objective of this study was to determine if a subterranean rodent that forages on aboveground plant parts, Ctenomys talarum, is able to detect the presence of odorous chemicals released by the plants into the soil to guide its excavation, as has been shown for subterranean species that forage on roots and bulbs. Individuals were introduced into an artificial Y‐maze, whose arms were filled with sandy soil in which plants either had or had not been growing (control). Digging activity exhibited in each soil type was recorded. The following plant species, present in the natural habitat of C. talarum, were used: Panicum racemosum (three different concentrations), Cortadeira sp., Margiricarpus pignatus, Ambrosia sp. and Raphanus sativus. The individuals allocated more time to excavation in the soil in which vegetation had been growing in comparison with the control for the five analysed plant species, except in the case of P. racemosum at its lowest concentration. The number of individuals that completed excavation in the tube‐containing soil in which plants had been growing was significantly higher than in the tube‐containing soil in which no plants had been growing, with the exception of P. racemosum at its lowest concentration and R. sativus. The results of the present study suggest that C. talarum have the ability to use olfaction to orient their digging while foraging even though members of this species consume mainly aboveground plant parts.