Coefficients of digestibility and nutritional values of geophytes and tubers eaten by southern African mole-rats (Rodentia: Bathyergidae)

Subterranean rodents have high energy requirements when they are excavating their burrows. This study investigates the energy available to, and the efficiency with which it can be extracted by, four species of bathyergid mole-rats fed natural diets ranging from the underground storage organs of geophytes to grass roots and leaves.
The digestibility coefficients of geophytes ranged from 53% for the fibrous tuber of the gemsbok cucumber to 95–7% for corms and bulbs. One species, Bathyergus suillus whose diet consists of over 80% grass, had a digestibility coefficient of 87% on an all grass diet.
All species had similar coefficients of digestibility of > 90% when fed on a uniform diet of sweet potato.
Bulbs and corms had a low fibre content (3–3–4%), high calorific value (15–16kJ/g) and high digestibility coefficients (95–7–96%) and on this diet the mole-rats maintained their body mass. Food of lower digestibility tended to have a high fibre content (8–2–45%) and, with the exception of B. suillus , although the mole-rats consumed a greater quantity of food, they lost mass. The sweet potato had a low fibre content (4–1 %) but was energetically very similar to bulbs and corms (15–5 kJ/g).
Geophytes which have low fibre contents are generally small (1–20 g), whereas geophytes with high fibre contents are much larger (30–2000 g) and often occur in more arid zones.     

Foraging in the subterranean social highveld mole-rat (Cryptomys hottentotus pretoriae): an investigation into mass-dependent geophyte use and foraging patterns

The foraging behaviour of captive colonies of the highveld mole-rat Cryptomys hottentotus pretoriae was investigated in an artificial soil-filled burrow system provided with four trays (patches) that varied in geophyte density and mass. An initial trial involving empty trays (only soil) revealed that there was no preference for any specific tray. There were no statistically significant preferences for excavating in any of the patches of different geophyte density. No preferences were evident for excavation in patches containing geophytes of different mass classes. Empty patches seemed to be preferred over patches containing geophytes when combinations of geophyte density/mass were investigated. The duration of handling and the rate of consumption of geophytes were recorded for 23 individuals of two mass classes. Handling time of geophytes was not related to mole-rat sex, but was strongly linked to mole-rat mass class. Handling time of geophytes was related to geophyte mass class. Small geophytes were less profitable to consume. These findings are considered in light of optimal foraging theory and the situation in the field. It was concluded that the mole-rats generally followed the qualitative predictions of optimal foraging theory, although falling short of being energy maximizers.     

The magnetic compass orientation of the burrows of the Damara mole-rat Cryptomys damarensis (Bathyergidae)

In response to reports claiming that part of the ability of mole-rats (Bathyergidae) to orientate with respect to the geomagnetic field involves orientation of their burrow systems in a southward direction, we measured the orientation of burrows of the Damara mole-rat, Cryptomys damarensis , in the Kalahari Desert. It was found that burrow orientation was not significantly different from that expected for a random distribution of compass orientations.     

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.     

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.     

Behaviour and social organization in a colony of the Damaraland mole-rat Cryptomys damarensis

This paper reports qualitative and quantitative behavioural studies of the social mole-rat C. damarensis . The behaviours were subdivided into four major categories: nest, maintenance, interactive and auto-behaviours. Social interactions and behavioural types were analysed by sex, age category and caste. The nest area is a focal point in the burrow system with individuals spending more than 70% of the day there. Many of the interactive behaviours occurred there and the mole-rats showed definite preferences with regard to which individuals they rested next to or avoided.
The Damaraland mole-rat colonies show some division of labour, into frequent and infrequent worker groups; these castes differed significantly in the amount of both digging and soil movement undertaken.     

Colony size and structure, activity patterns and foraging behaviour of a colony of the social mole-rat Cryptomys damarensis (Bathyergidae)

A complete colony of 20 Cryptomys damarensis was trapped in the Kalahari Gemsbok National Park, South Africa. The colony consisted of 15 males and five females and formed three distinct size classes, two of which were presumed to represent worker (smallest mole-rats) and soldier (largest males) castes. Body masses ranged from 86–197 g, with the dominant male being the largest mole-rat. The activity patterns of five animals in the colony were monitored by radiotelemetry. Activity patterns were not influenced by diel fluctuations in burrow temperature and photoperiod. It was suggested that the chance of hyperthemia influenced activity, such that in order to maximize the daily distance burrowed (foraging efficiency), the mole-rats had to engage in frequent (5.58 times. day^-1), but short ( c. 60 min) foraging bouts.     

Experimental manipulation of predation risk and food quality: effect on grazing behaviour in a central-place foraging herbivore

The relative importance of predation risk and food quality on spatial grazing pressure and activity patterns was tested in a central-place foraging herbivore: the European rabbit. Rabbits grazed less with increasing distance from their burrows into adjacent grassland, thereby creating a gradient of increasing vegetation height and plant biomass and decreasing plant nutrient concentration. When nitrogen concentration was experimentally increased by 150% through fertilizing and mowing, rabbits visited these plots four times more frequently than the untreated control plots. Addition of predator scent (mink pellets) did not result in different patch use by rabbits. The combined addition of fertilizer and mink pellets had the same effect as addition of fertilizer alone. However, the mink pellets changed the temporal activity pattern of rabbits as measured with infrared detectors. Rabbits were predominantly nocturnal but shifted their activities to the day when mink pellets were added, resulting in equal activities during night and day. We conclude that rabbits are sensitive to perceived predation risk, but that this does not influence their spatial grazing pressure. A selection for the highest food quality on the other hand can explain the observed natural rabbit grazing gradient. Food quality was highest close to the burrows, therefore rabbits selecting for high quality food should forage most intensely close to the burrows and only move further away for higher quality items or when the vegetation close to their burrows is depleted. Through intensive grazing close to the burrows rabbits facilitated for themselves either through stimulating fresh protein rich re-growth or the return of nutrients through faeces or both. This is in contrast with central-place foraging theory where intense feeding close to the burrow is assumed to lead to reduced food resources.     

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.     

Eusociality in African mole-rats: new insights from patterns of genetic relatedness in the Damaraland mole-rat (Cryptomys damarensis)

After the discovery of eusociality in the naked mole-rat, it was proposed that inbreeding and high colony relatedness in this species were the major underlying factors driving cooperative breeding in African molerats. By contrast, field and laboratory studies of the eusocial Damaraland mole-rat (Cryptomys damarensis) have raised the possibility that this species is an obligate outbreeder, although the build-up of inbreeding over several generations could still occur. Using microsatellite markers, we show that most breeding pairs in wild colonies of the Damaraland mole-rat are indeed unrelated (R = 0.02 +/- 0.04) and that mean colony relatedness (R = 0.46 +/- 0.01), determined across 15 colonies from three separate populations, is little more than half that previously identified in naked mole-rats. This finding demonstrates that normal familial levels of relatedness are sufficient for the occurrence of eusociality in mammals. Variation in the mean colony relatedness among populations provides support both for the central role played by ecological constraints in cooperative breeding and for the suggestion that inbreeding in naked mole-rats is a response to extreme constraints on dispersal. Approaches that determine the relative importance of an array of extrinsic factors in driving social evolution in African mole-rats are now required.     

The colony structure and dominance hierarchy of the Damaraland mole-rat, Cryptomys damarensis (Rodentia: Bathyergidae), from Namibia

Cryptomys damarensis is one of the few subterranean rodents which is social. This species is found in the semi-arid regions of southwestern and central Africa. The Damaraland mole-rat occurs in coloniesof up to 25 individuals, in which reproduction is limited to one or two of the largest males and the largest female in the colony.
The mean colony size is 18 mole-rats ( n =6 colonies). The mean colony biomass is 2.32 kg and the sex ratio is female biased (0.71–0.78). The number of mole-rats in each colony, the mean body mass and the sex ratio are described for six field-captured colonies, three of which were captured in their entirety.
The dominance hierarchy of two colonies of C. damarensis was found to be linear with a value of between 0.94 and 1.00 calculated from Landau's linearity index. Dominance was found to be related to gender, with the males more dominant than females. The reproductive individuals are the dominant animals within each respective gender. The non-reproductive females rank lowest in the hierarchy.     

An assessment of the breeding range,colony sizes and population of the Westland petrel (Procellaria westlandica)

Abstract

The Westland petrel (Procellaria westlandica) is an endemic New Zealand species and one of the very few burrowing seabird species still breeding on mainland New Zealand. It nests only on a series of coastal ridgelines near to Punakaiki on the West Coast of the South Island. Between 2002 and 2005, surveys were undertaken at 28 of the 29 known colonies. The area occupied by the colonies was 73 ha; most colonies had fewer than 50 burrows, but six colonies had 201–500 burrows and four colonies had more than 1000 burrows. We find that the current breeding range of Westland petrel and the location of individual colonies are similar to those reported in both the 1950s and 1970s. Based on total burrow counts at 28 colonies and burrow occupancy rates determined by annual monitoring, the annual breeding population is estimated to be between 2954 and 5137 breeding pairs.     

Fibre utilization by Kalahari dwelling subterranean Damara mole-rats (Cryptomys damarensis) when fed their natural diet of gemsbok cucumber tubers (Acanthosicyos naudinianus)

Kalahari dwelling Damara mole-rats (Cryptomys damarensis) naturally feed on a high fibre diet of underground gemsbok cucumber tubers, Acanthosicyos naudinianus. We investigated the degree of fibre utilization and fermentation on this diet by measuring caecal characteristics (namely temperature, pH and weight) and in vitro rates of gas and short chain fatty acid (SCFA) production in these underground dwelling hind-gut fermentors. Rectal temperatures (33.8±0.6°C) were consistently higher than caecal temperatures (33.3 ± 0.6°C). Furthermore, a 0.8°C gradient of temperatures existed within the caecum, with the lowest temperature occurring in the corpus caeci. Both rates of gas production (4.74 ± 0.6 ml/g dry matter/hr) and SCFA production (266.80 ± 9.251 /i mol/caecum per hr) were high, with proportionately more acetic acid produced than any other SCFA. Nevertheless, the initial concentrations of SCFAs present in the caecum were low (52 ±17 mM) implying a rapid rate of absorption of these SCFAs. The high rates of fermentation provide a considerable amount of energy that would otherwise be trapped in fibre and thus unavailable to the animal. This highly efficient caecal fermentation enables the Damara mole-rat to maximally exploit the underground food resources in the arid-zone ecotope.     

On foraging,recruitment systems and optimum number of scouts in eusocial colonies

Summary A numerical model of an eusocial colony foraging for food showed that, for each set of values of resource density, resource size and recruitment system employed, a given optimal proportion of scouts in the colony maximize the amount of resources retrieved by a colony during a fixed period. The model predicts that ants using mass recruitment systems should have larger colonies with small foragers, and should forage on large food sources. Retrieval of small food sources by small colonies is best achieved with large workers using individual foraging strategies. For mass foragers, several food sources are best retrieved using democratic decision-making systems in recruitment, whereas for very large food sources at very low mean food patch density, autocratic decision-making systems are optimal. Some of the experimental evidence available is discussed in the light of these findings, as they confirm the prediction that large colonies with small workers have mass recruitment systems, whereas workers of small colonies with large workers are generally lone foragers.     

Social cues elicit sexual behavior in subordinate Damaraland mole-rats independent of gonadal status

Damaraland mole-rats (Fukomys damarensis) are among a small number of eusocial mammals. Eusociality is a social system where only a few individuals within a colony engage in direct reproduction, while remaining subordinate members are non-breeders and support reproductive efforts of breeding individuals. Inbreeding avoidance precludes mating between subordinate siblings and between offspring and parents. Interestingly, non-breeders readily attempt to mate with unrelated opposite-sex individuals. This is unusual since the non-breeding females do not attain puberty while in their natal colony. Based on this finding, the present study investigated the role of the gonads in the regulation of mating behaviors in this species and identified the mechanism of inbreeding avoidance. Gonadal-intact and gonadectomized non-breeders from different colonies were removed from their colonies and tested for the expression of sexual behavior. Results indicated that gonadal status had only minor effects on the expression of sexual behavior in either males or females. In a second experiment, sexual behaviors were absent between opposite-sex siblings so long as they had frequent contact with each other; however, following 5 weeks of separation, sexual behavior between these siblings was robustly expressed. Thus, Damaraland mole-rats avoid establishing mating relationships with familiar individuals but will readily mate with unfamiliar individuals of the opposite sex, with genetic relatedness apparently playing little role. The initiation of sexual behavior in Damaraland mole-rats does not require the presence of the gonads, but does require that the members of the pair have not been in contact with one another for at least several weeks.     

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.     

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.     

Aggregation in a Desert Tenebrionid Beetle: A Cost/Benefit Analysis

Parastizopus armaticeps (Coleoptera: Tenebrionidae), a nocturnal fossorial detritivore inhabiting southern Kalahari dunes, aggregates in burrows during the day. Group size increases during drought but 25% of beetles are still found alone or in pairs. During drought, beetles from large groups leave burrows after sunset synchronously and carlier than pairs and single animals and earlier than beetles of any group size after rain. Detritus from the beetles' major foodplant is scarce and food competition high. Beetles emerging early preferentially select and carry high-quality transportable items into burrows to eat (forage); late-emerging ones feed on the low-quality large twigs on the surface. Foraging is shown to be a strategy to secure food items against surface competitors, not one to reduce body water loss during surface exposure. The costs and benefits of group vs. solitary lifestyles and alternate hypotheses for early and synchronous emergence were tested experimentally. Grouped beetles had lower body water loss rates but, due to competition with burrow mates, higher feeding costs than single ones. It is hunger that advances and thus synchronizes emergence time, not social facilitation. Field data support a model predicting that, for maximal benefits, beetles should alternate between solitary and group life at optimal time intervals.     

Small- and large-scale effect of the SW Atlantic burrowing crab Chasmagnathus granulatus on habitat use by migratory shorebirds

Here we address the question of whether the presence of the burrowing crabs Chasmagnathus granulatus affects small- and large-scale habitat use by migrant shorebirds. This crab is the dominant species in soft bare sediments and vegetated intertidal areas along the SW Atlantic estuaries (southern Brazil 28°S to the northern Argentinean Patagonia 42°S). They generate very extensive burrow beds in soft bottom intertidal areas. Our information shows that this burrowing crab affects the small-scale habitat use by shorebirds, given that shorebirds never walk through the funnel-shaped entrances of burrows. Given that crab burrow entrances occupy up to 40% of the intertidal area, there is a large decrease of available shorebird habitat in crab beds, restricting their activity to the spaces between the burrows. The southern migratory shorebird Charadrius falklandicus maximize the use of these areas by foraging closer to the burrows than the other bird species. Neotropical migrants, such as Calidris fuscicollis, Pluvialis squatarola and Tringa melanoleuca, used foraging paths that tended to maximize the distance from burrows, especially the distance to larger burrows. A field experiment showed that this was not necessarily due to a decrease in the availability of polychaetes near the crab burrows. A combination of landscape measurements and satellite images showed that crab beds covered up to 40% of the intertidal area of the Mar Chiquita coastal lagoon (37°40′S, Argentina), and nearly 100% of the intertidal area of the Bahia Blanca estuary (38°48′-39°25′S, Argentina). These two estuaries are located along the migratory flyway of Neotropical migratory shorebirds, but the Bahia Blanca estuary (area∼110,000 ha) shows a much lower shorebird diversity than Mar Chiquita (area∼4500 ha). The most common species in Bahia Blanca is the two-banded plover C. falklandicus, the species least affected by crabs at Mar Chiquita and which prefers to use high-density crab areas as foraging sites. The oystercatcher Haematopus palliatus was also most abundant in high-density crab areas, but they used these areas for resting. The abundances of preys varied during the study period and between the crab density areas, indicating that the use of these areas by birds is independent of crab density. However, burrowing crabs affect the depth distribution of polychaete and thus their availability to shorebirds. We suggest that this shorebirds-burrowing organism interaction could be generalized for other intertidal estuarine habitats.     

Burrow architecture,family composition and habitat characteristics of the largest social African mole-rat: the giant mole-rat constructs really giant burrow systems

Among African mole-rats, the giant mole-rat Fukomys mechowii is the largest social species. Despite several attempts to study a free-living population, information on its biology from natural habitats is very scarce. We mapped two neighbouring burrow systems of the giant mole-rat in a miombo woodland in Zambia. We provide information on the size and kin structure of the respective mole-rat families, architecture of their burrow systems, and characteristics of the food supply and soil around the two mapped and additional ten burrow systems. Both uncovered burrow systems were very large (total lengths, 2,245 and 743 m), making them the largest burrow systems ever mapped. Food resources around the additional ten burrow systems had a clumped distribution (standardized Morisita index of dispersion = 0.526), but a relatively high biomass (298 ± 455 g m⁻²). This, together with favourable soil conditions even in the advanced dry season (cone resistance, 328 ± 50 N m⁻²; soil density, 1.36 ± 0.06 g cm⁻³) indicates relatively hospitable ecological conditions. Both food supply and soil conditions were comparable with the conditions found in a miombo habitat of the solitary silvery mole-rat in Malawi. This suggests that there are no ecological constraints which would preclude the solitary life of a subterranean herbivore from the examined habitat. Microsatellite analysis supported the assumption that giant mole-rats live in monogamous multigenerational families with only one breeding pair of non-related animals and their offspring. The mean family size is consistent with previous findings on this species and comparable to that found in other Fukomys species studied thus far.