Absentee Mothering – Not So Absent? Responses of European Rabbit (Oryctolagus cuniculus) Mothers to Pup Distress Calls

Longer‐range acoustic parent‐offspring communication is widespread, but might be absent in species in which young are hidden in burrows during the mother's absence. The European rabbit (Oryctolagus cuniculus) is such a species, with unusually limited maternal care largely restricted to a short daily nursing bout. Based, however, on evidence of frequent infanticide in this species, we hypothesize that rabbits possess a mechanism promoting a maternal response to pup distress calls. We conducted playback experiments with distress calls of pre‐weaning pups played next to the breeding burrows of mothers in a field enclosure (i.e. next to the burrows where mothers give birth and raise their young). Calls were played shortly after pups were born (T1) when infanticide risk is maximal, and shortly before the pups start dispersing from the breeding burrow (T2). A high proportion of mothers (60.6%) responded to pup calls by rapidly returning to their breeding burrow and 40% of them investigated the area around the entrance. Return responses to the playback of pup calls did not differ between mothers during T1 and T2. Thus, our results confirm that rabbit mothers respond rapidly to pup distress calls and that this responsiveness may adaptively serve to repel potentially infanticidal females.     

Elevated predation risk changes mating behaviour and courtship in a fiddler crab

The fiddler crab, Uca beebei, lives in individually defended burrows, in mixed-sex colonies on intertidal mud flats. Avian predation is common, especially of crabs unable to escape into burrows. Mating pairs form in two ways. Females either mate on the surface at their burrow entrance (''surface mating'') or leave their own burrow and sequentially enter and leave (''sample'') courting males'' burrows, before staying in one to mate underground (''burrow mating''). We tested whether perceived predation risk affects the relative frequency of these mating modes. We first observed mating under natural levels of predation during one biweekly, semi-lunar cycle. We then experimentally increased the perceived predation risk by attracting grackles (Quiscalus mexicanus) to each half of the study site in two successive biweekly cycles. In each experimental cycle, crabs were significantly less likely to mate on the side with more birds. Moreover, on the side with elevated predation risk, the number of females leaving burrows to sample was greatly reduced relative to the number of females that surface-mated. Males waved less and built fewer mud pillars, which attract females, when birds were present. We discuss several plausible proximate explanations for these results and the effect of changes in predation regime on sexual selection.     

Burrow webs: Clawing the surface of interactions with burrows excavated by American badgers

Ecosystem engineers are organisms that influence their environment, which includes alterations leading to habitat provisioning for other species. Perhaps the most well‐examined guild of species provisioning habitat for other species is tree cavity excavators or woodpeckers (Picidae). Many studies have examined the suite of secondary cavity users that rely on woodpeckers, and how the ecological network of secondary users, collectively referred to as the nest web, changes across communities. Despite similar habitat provisioning processes, fewer studies have assessed the suite of species associated with burrowers providing access to subterranean habitat. Here, we begin to characterize the burrow web provisioned by American badgers (Taxidea taxus) and evaluate the diversity and frequency of species interactions we detected at abandoned badger burrows in Wyoming, USA. We deployed camera traps at 23 badger burrows and identified interactions with the burrow by birds, mammals, and reptiles. Overall, we discovered 31 other species utilizing badger burrows, consisting of 12 mammals, 18 birds, and 1 reptile. Mammals, other than American badgers themselves and other fossorial species such as ground squirrels (Urocitellus sp.), frequently using burrows included mice (Peromyscus sp.), long‐tailed weasel (Mustela frenata), pygmy rabbit (Brachylagus idahoensis), and desert cottontail (Sylvilagus audubonii). Of the 18 bird species detected, most accounted for <5% of overall detections, besides chipping sparrows (Spizella passerina) at 7.2%–11.5% of detections. The most common category of detection by bird species was foraging, contrary to mammals, which used the burrow frequently and were commonly observed entering and exiting the burrow. This work provides additional context on the ecological role of American badgers within their environment. More broadly, this work scratches the surface of many remaining questions to explore with the aim of advancing our understandings about burrow webs across the diversity of burrowing species and the communities in which they occur.     

How Do Carpenter Bees Recognize the Entrance of their Nests?: An Experimental Investigation in a Natural Habitat1

Carpenter bee females (genus Xylocopa) enter only their own nests. They can be found in the same burrow over a period of months, regardless of whether they arc living singly or aggregated in close vicinity. The aim of this study was to investigate and to clarify which sensory systems are responsible for the recognition of the nest. Females of three different African species were tested in their natural habitat. Following alteration in the immediate visual surroundings of either the entrance or the entire dwelling stem, bees behaved as they did in undisturbed situations and entered their nests. Furthermore, a relative dislocation of the entrance within the dwelling stem or a displacement of the whole stem within a restricted range had no effect on the recognition of the burrow. When the entrance was plugged by foam rubber the bees landed only after extended searching. When the entrance was closed by a strip of scotch tape the bees searched for several minutes, departed, then returned and searched again with the same result. The bees showed no indications that they recognized their burrow. These results strongly support the conclusion that the bees recognized their burrows primarily or exclusively by olfactory cues, and, furthermore, that individual odors were involved. The possible contributions of different sensory systems for the recognition of the nest and the possible biological function of the use of olfactory cues are discussed.     

Influence of sediment characteristics on density and distribution of Ocypodoid crab burrows (superfamily: Ocypodoidea) along the coastal areas of Pakistan

Crabs belong to the superfamily Ocypodoidea are a significant component of benthic fauna and considered as ecosystem engineers because of their dynamic role as an active burrower in mangrove and estuarine environment. The current investigation was to evaluate the crab burrow density, diameter and total area of burrow opening along the coast of Pakistan. The variations in burrow properties and their relation to sediment characteristics were also evaluated to recognize the most influencing variables of sediments that effects on crab burrows. All crab burrow and sediment characteristics differed significantly (p < 0.05) among the monitoring sites. Regression analysis showed that crab density was significantly correlated with burrow density (P < 0.001). Moreover, burrow density was noticed significantly greater (p < 0.05) than crab density. Pearson correlation analysis reveals that moisture, porosity, organics, sand and mean grain size observed as most influencing the features of sediment to determine the ecological functioning of crab burrows in mangrove and mudflats of Pakistan.     

Habitat selection by the Spiny-tailed lizard (Uromastyx aegyptia): A view from spatial analysis

Many factors affect the habitat selection for animal species, which in turn may greatly affect their distribution in different ecosystems. Understanding the processes that affect habitat selection is also critical for guiding and managing conservation initiatives. Our study aimed to assess the habitat selection by free-ranging Spiny-tailed lizard (Uromastyx aegyptia) by analyzing a geospatial data connecting its burrow parameters to different habitat characteristics within selected sites in Hail region, Saudi Arabia. We examined evidence and patterns of significant spatial clustering for (366) active burrows by linking their parameters (burrow entrance size, burrow entrance width and burrow entrance height), their reference geographical locations and, two habitat characteristics defined by soil type and vegetation cover. The objective of the analysis was to increase the understanding on the burrows aggregation process in the space and, to describe its possible relation to other spatial habitat configurations. Analysis of distances based on the Nearest Neighbor Index (NNI) and hotspots detection in Nearest neighbor hierarchical clustering (Nnh) suggested twelve (12) spatial clusters located within the study area. In addition, a spatial ordinary least square (OLS) and Poisson regression models revealed significant effects of soil type and vegetation cover on burrow parameters (OLS, p < 0.05; Poisson, p < 0.001), which indicate a strong association between burrows parameters and habitats characteristics. Findings from the study also suggest that other factors such as elevations, highways, and human settlement concentration spots could possibly play a major role in defining burrow spatial aggregation and furthermore have a significant impact on habitat selection.     

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.     

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.     

Review of bees as guests in termite nests,with a new record of the communal bee, <Emphasis Type="Italic">Gaesochira obscura</Emphasis> (Smith, 1879) (Hymenoptera,Apidae), in nests of <Emphasis Type="Italic">Anoplotermes banksi</Emphasis> Emerson, 1925 (Isoptera,Termitidae, Apicotermitinae)

Associations between bees and termites are documented infrequently, but records are available for bee species ranging in behavior from solitary to highly eusocial. The subtribe Meliponina (stingless bees) is the most common bee group reported in termite nests, and, for some species, the occupancy of termite nests may be obligatory. The records of solitary bees nesting within termite nests include species of the tribes Emphorini, Centridini, Megachilini, and Paracolletini. Most of these bees can probably nest in other substrates, and their relationships with termite nests are presumably opportunistic. This study provides a review of published records of bees as guests in termite nests, and also describes the aggregation of nests of Gaesochira obscura within one nest of Anoplotermes banksi in Brazilian Amazonia. One termite nest contained at least ten nest entrances of G. obscura, with burrows 4–6 mm in diameter and about 10 cm long. Each burrow ended in brood cells in different stages of food provisioning and larval development. As commonly reported for other associations of this nature, there was no connection between the tunnels of bees and those of termites. This record adds important data on the biology of A. banksi. Because this is a single record, it is impossible to classify G. obscura either as a termitophile or termitariophile; this species may be opportunistic in relation to nesting substrate.     

Model of burrow selection predicts pattern of burrow switching by Leach's Storm‐Petrels

Patterns of nest site selection exhibited at the scale of a population should result from initial preferences of individuals occupying nest sites as well as preferences exhibited by individuals moving between nest sites. We tested whether nest‐site preferences measured at the population scale were predictive of patterns of burrow switching by Leach's Storm‐Petrels (Oceanodroma leucorhoa), a long‐lived seabird that nests in underground burrows. Breeding pairs generally choose from the pool of available existing burrows rather than constructing new burrows, and a portion of the burrows in a colony remains unused in any breeding season. We quantified burrow preference at a colony on Kent Island, New Brunswick, over four breeding seasons. We used a classification and regression tree analysis to build a predictive model of nest‐site selection. Preferentially occupied burrows were drier, longer, had larger nest chambers, and were in areas of higher burrow density. To measure preferences during burrow switching, we tracked individuals that switched burrows, comparing characteristics of the burrows in which these birds were originally found to those they inhabited at the end of the study period. Characteristics preferred by switching individuals were a subset of those observed at the scale of the population; individuals moved to burrows that were drier, longer, and had larger nest chambers. Our results show how preferences of individuals that move between nest sites contribute to nest site preferences exhibited at the population scales commonly tested.     

Effects of group size on nest attendance in the communally breeding colonial tuco-tuco

Communal nesting is generally assumed to be adaptive, meaning that it confers a fitness advantage on the individuals that share a nest site. This advantage may accrue directly to adults, or it may affect adult fitness through gains in offspring survival. In particular, survival of juveniles reared in communal groups may be greater because adults are present in the nest more often to provide care to young. To test the hypothesis that communal nesting is associated with increased adult presence in the nest, we used radiotelemetry to examine patterns of adult nest attendance as a function of group size for free-living colonial tuco-tucos (Ctenomys sociabilis). Burrow systems of this social, subterranean rodent are inhabited by 1–6 adult females and, in some cases, a single adult male. Data obtained from residents of 26 burrow systems monitored during 1996–2000 indicated that the percentage of time that the nest was unattended (no adult present) did not vary predictably with date or time of day during the period between the birth and weaning of young. The percentage of time that the nest was unattended, however, decreased significantly as the number of adults per burrow system increased. This difference was most evident when the percentage of time that the nest was unattended was compared for lone females versus multi-adult groups. We suggest that increased nest attendance has important implications for the survival of juveniles reared in multi-adult burrow systems but that this effect may be confounded by the fitness consequences of other costs and benefits associated with communal nesting in this species.     

Comparative Burrow Architectures of Resident Fiddler Crabs (Ocypodidae) in Indian Sundarban Mangroves to Assess Their Suitability as Bioturbating Agents

Excavation of burrows by fiddler crabs (genus Uca) is an important component in mangrove ecosystem functioning. This bioturbation activity can be measured by analysing the burrow architecture of these crabs. The aim of the present study is to describe and evaluate inter specific differences in the burrow morphologies of four species of fiddler crabs (Uca rosea, Uca triangularis, Uca dussumieri and Uca vocans) using polyester resin casts of the burrows. For each of the species, sex and carapace width (CW; mm) were determined for all the individuals. Three burrow morphological characters viz. burrow diameter (BD; mm), total burrow depth (TBD; mm) and burrow volume (BV; cm³) were considered during the study. Density of each species throughout the year was also assessed. For all the species BD and BV were higher in case of males compared to the females and they showed significant positive correlation with the CW of the burrow inhabitants. The amount of sediment excavated by each crab was evaluated in terms of BV. Among all the studied species, U. rosea was established as the most potent bioturbative candidate in the studied mangrove due to their greater density and moderate ability to excavate burrow.     

Nesting and biology of Jucancistrocerus caspicus (hymenoptera,vespidae, eumeninae)

Females of Jucancistrocerus caspicus nest in dense clay ground on the vertical surface of cliffs. The nests contain 1–9 cells (on average 2.8) and have a linear-branched construction. Females surmount the entrance of the burrow with a curved chimney which has a laced structure. The cells are positioned vertically in the main burrow and obliquely or horizontally in the lateral tunnels; the cells in a row are separated with double partitions. The size of the cells is 7–9 × 4–4.5 mm, the diameter of the nest burrow is 4 mm. The egg is laid before provisioning and is attached to the cell ceiling with a filament. Females hunt for weevil larvae and store 23–33 larvae (on average 27.8) in each cell. The species is univoltine, with prepupae hibernating in their cocoons. The nests are parasitized by the cuckoo wasps Chrysis rutilans which cause 11.5% of brood mortality. Adult wasps are killed by the spiders Pholcus sp. living near the nests.     

Predatory behaviour in the ant-like wasp Methocha stygia (Say) (Hymenoptera: Tiphiidae)

An examination of prey-orienting behaviour of Methocha stygia has demonstrated that adult females regularly transport prey (Cicindela larvae) when they are removed from their burrows. On a single occasion a wasp was also observed to utilize a naturally occurring depression in the soil as a nest site, instead of depending on the prey's own burrow. This flexibility in response might be a carryover of the behaviour that surely existed in the ancestors of the methochine wasps, insects which were close to if not identical with the ancestors of the ants. When Methocha females were crowded together with prey in the laboratory, frequent avoidance and aggressive behaviours were noted, but no sign of any form of co-operation.     

Evolutionarily stable nesting strategy in a digger wasp.

Two alternative “strategies” will not coexist in a population unless on average they are equally successful. The most likely way for such an equilibrium to be maintained is through something equivalent to frequency-dependent selection. Females of the digger wasp Sphex ichneumoneus (Sphecidae) nest in underground burrows. They usually dig and provision these by themselves but occasionally a nest is jointly occupied. The two wasps fight whenever they meet and in the end only one of the two females lays an egg in the shared nest. Two models based on the theory of mixed evolutionarily stable strategies were developed and tested on comprehensive field data from two North American populations of these wasps. The first model proposes two strategies called founding and joining. Founders start burrows alone, but they are more successful when they are joined by a joiner. At equilibrium founders and joiners are equally successful, which amounts to an amicable, sharing relationship. The predictions of this amicable model are decisively rejected by the data. The second model proposes two strategies called digging and entering. Diggers dig their own burrows but they often have to abandon these burrows because of temporary unsuitability. Enterers move in later, thereby exploiting abandoned burrows as a valuable resource. They do not distinguish an adandoned burrow from one that is still occupied. Therefore sharing of burrows arises as an unfortunate by product of selection for entering abandoned burrows, and Model 2 is not an amicable model. Its quantitative predictions are impressively fulfilled in one population, though not in another population. This is one of the only examples yet known of a mixed evolutionarily stable strategy in nature. Yet the word strategy itself can confuse, and this paper tries the experiment of substituting “decision”, defined as a moment at which the animal commits future time to a course of action.     

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.     

Obligate crayfish burrow use and core habitat requirements of crawfish frogs

Crawfish frogs (Lithobates areolatus) have experienced declines across large portions of their former range. These declines are out of proportion to syntopic wetland-breeding amphibian species, suggesting losses are resulting from unfavorable aspects of non-breeding upland habitat. Crawfish frogs get their common name from their affinity for crayfish burrows, although the strength of this relationship has never been formally assessed. We used radiotelemetry to address 4 questions related to upland burrow dwelling in crawfish frogs: 1) what burrow types are used and how do they function to affect crawfish frog survivorship; 2) what are the physical characteristics and habitat associations of crawfish frog burrows; 3) what are the home range sizes of crawfish frogs when burrow dwelling; and 4) where are crawfish frog burrows situated with respect to breeding wetlands? We tracked crawfish frogs to 34 burrows, discovered another 7 occupied burrows, and therefore report on 41 burrows. Crawfish frogs exclusively occupied crayfish burrows as primary burrows, which they inhabited for an average of 10.5 months of the year. With one exception, crawfish frogs also used crayfish burrows as secondary burrows—temporary retreats occupied while exhibiting breeding migrations or ranging forays. Burrows were exclusively located in grassland habitats, although crawfish frogs migrated through narrow woodlands and across gravel roads to reach distant grassland primary burrow sites. Home range estimates while inhabiting burrows were 0.05 m² (the area of the burrow entrance plus the associated feeding platform) or 0.01 m³ (the estimated volume of their burrow). Crawfish frog burrows were located at distances up to 1,020 m from their breeding wetlands. To protect crawfish frog populations, we recommend a buffer (core habitat plus terrestrial buffer) of at least 1.2 km around each breeding wetland. Within this buffer, at least 3 critical habitat elements must be present: 1) extensive grasslands maintained by prescribed burning and/or logging, 2) an adequate number of upland crayfish burrows, and 3) no soil disturbance of the sort that would destroy crayfish burrow integrity. © 2012 The Wildlife Society.     

Giving your daughters the edge: bequeathing reproductive dominance in a primitively social bee

The fitness associated with behavioural strategies is usually estimated in terms of offspring number and size. However, in group-living animals the reproductive value of offspring may also depend on their social rank. We show here that in an allodapine bee Exoneura robusta, dominant mothers can behaviourally influence their daughters'' reproductive rank by controlling insemination of other potential mothers. In E. robusta, group living is near mandatory and reproductive dominance among female nestmates is determined by order of adult emergence. Nests are single, undivided burrows and the dominant female assumes a guarding position closest to the nest entrance. We show that before the egg-laying period, subordinate females who have been absent from the nest are ''screened'' by the reproductive guard upon attempted re-entry. Those who have been in contact with foreign males are less likely to be granted access back into the nest than those who have been in contact with foreign females or with no bees at all. We argue that by controlling insemination patterns of their nestmates, dominant females ensure that their own daughters eclose first and are therefore more likely to assume dominance in the next generation. This presents a situation where dominance is bequeathed to daughters by behavioural means. The ability of mothers to influence social hierarchies in subsequent generations introduces a fitness component additional to the number and size of offspring produced.     

Estimating and Correcting for Bias in Population Assessments of Sooty Shearwaters

Abstract: We investigated the precision and accuracy of an infrared burrowscope for detecting sooty shearwater (Pufffinus griseus) chicks at 13 plots from 3 islands in southern New Zealand in 2003. We partially excavated burrows systems to reveal the entire burrow contents after 2 teams of observers had prospected all burrow entrances. Accuracy was similar between islands and observer teams at approximately 85%. The majority of the inaccuracy stemmed from failure to detect some chicks. Logistic regression modeling identified 4 burrow characteristics occurring between the entrance and the nest-site that influenced detection of burrow occupants. Detection was lower at nest-sites further from burrow entrances, in burrows with a high rate of burrow division, and in burrows with a high level of curvature. There was a positive relationship between the interaction of rate of division and curvature and detection of chicks. Distance from the burrow entrance was the only parameter that could be reliably used as a predictor of detection rate, so a reduced model containing only this variable was constructed to correct for burrowscope bias. The correction factor performed well on The Snares and Bench Island where predicted bias was very similar to observed levels (within 5%), but bias was overestimated on Putauhinu by up to 19.1%. Consistent bias, lack of damage to burrows from excavation, and the successful application of a correction factor all indicate the value of further testing burrowscope accuracy on other burrow-nesting seabird species.     

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.