Evidence for contrasting roles for prolactin in eusocial naked mole-rats, Heterocephalus glaber and Damaraland mole-rats, Fukomys damarensis.

Elevated prolactin (PRL) has been associated with the expression of social and cooperative behaviours in a number of vertebrate species, as well as suppression of reproduction. As social mole-rats exhibit both of these traits, PRL is a prime candidate in mediating their social phenotype. While naked and Damaraland mole-rats (NMRs and DMRs) have evolved eusociality independently within their family, both species exhibit an extreme skew in lifetime reproductive success, with breeding restricted to a single female and one or two males. Non-breeding NMRs of both sexes are physiologically inhibited from reproducing, while in DMRs only the non-breeding females are physiologically suppressed. Newly emerging work has implicated the dopamine system and PRL as a component in socially induced reproductive suppression and eusociality in NMR, but the DMR remains unstudied in this context. To investigate evolutionary convergence in the role of PRL in shaping African mole-rat eusociality, we determined plasma PRL concentrations in breeders and non-breeders of both sexes, comparing DMRs with NMRs. Among samples from non-breeding NMRs 80% had detectable plasma PRL concentrations. As a benchmark, these often (37%) exceeding those considered clinically hyperprolactinaemic (25 ng ml-1) in humans: mean ± s.e.m.: 34.81 ± 5.87 ngml-1; range 0.00-330.30 ng ml-1 Conversely, 85% of non-breeding DMR samples had undetectable values and none had concentrations above 25 ng ml-1: 0.71 ± 0.38 ng ml-1; 0.00-23.87 ngml-1 Breeders in both species had the expected variance in plasma PRL concentrations as part of normal reproductive function, with lactating queens having significantly higher values. These results suggest that while elevated PRL in non-breeders is implicated in NMR eusociality, this may not be the case in DMRs, and suggests a lack of evolutionary convergence in the proximate control of the social phenotype in these mole-rats.

Molecular insight into patterns of colony composition and paternity in the common mole-rat Cryptomys hottentotus hottentotus.

We report the discovery of intraspecific variation in both colony composition and patterns of paternity in two populations of the social common mole-rat Cryptomys hottentotus hottentotus. These two populations represent the mesic and arid habitat extremes of the species' broad ecological range in South Africa. Until recently colonies of the common mole-rat were thought to consist of familial groups whereby all colony members were the offspring of a monogamous reproductive pair. The remaining colony members were thought to forego reproduction until both social and ecological conditions favoured dispersal and opportunities for independent outbreeding. Results from genetic assignment tests using microsatellite markers indicate that while colony composition is dominated by familial groups, colonies within both populations included both adult and subadult foreign conspecifics. Analysis of parentage reveals that the social organization of C. h. hottentotus is not that of strict monogamy; paternity of offspring was not assigned consistently to the largest, most dominant male within the colony. Moreover, a number of significantly smaller males were found to sire offspring, suggesting a sneak-mating strategy by subordinate within-colony males. Extra-colony extra-pair paternity (ECP) was also found to characterize C. h. hottentotus colonies, occurring with similar frequencies in both habitats. Both dominant established breeding males and subordinate males were identified as siring young in nonsource colonies. Furthermore, established breeding males were found to sire extra-colony young in the same season as siring young within their source colonies. We discuss the significance of these results within the context of the divergent ecological regimes characterizing the two sites and observe that our results revisit the accuracy of using behavioural and morphological characters, which have structured the basis of our understanding of the behavioural ecology of this species, as indicators of breeding status in mark-recapture studies.

Phylogeographical patterns of genetic divergence and speciation in African mole-rats (Family: Bathyergidae).

African mole-rats are subterranean Hystricomorph rodents, distributed widely throughout sub-Saharan Africa, and displaying a range of social and reproductive strategies from solitary dwelling to the 'insect-like' sociality of the naked mole-rat, Heterocephalus glaber. Both molecular systematic studies of Rodentia and the fossil record of bathyergids indicate an ancient origin for the family. This study uses an extensive molecular phylogeny and mitochondrial cytochrome b and 12s rRNA molecular clocks to examine in detail the divergence times, and patterns of speciation of the five extant genera in the context of rift valley formation in Africa. Based on a value of 40-48 million years ago (Myr) for the basal divergence of the family (Heterocephalus), we estimate divergence times of 32-40 Myr for Heliophobius, 20-26 Myr for Georychus/Bathyergus and 12-17 Myr for Cryptomys, the most speciose genus. While early divergences may have been independent of rifting, patterns of distribution of later lineages may have been influenced directly by physical barriers imposed by the formation of the Kenya and Western Rift, and indirectly by accompanying climatic and vegetative changes. Rates of chromosomal evolution and speciation appear to vary markedly within the family. In particular, the genus Cryptomys appears to have undergone an extensive radiation and shows the widest geographical distribution. Of the two distinct clades within this genus, one exhibits considerable karyotypic variation while the other does not, despite comparatively high levels of sequence divergence between some taxa. These different patterns of speciation observed both within the family and within the genus Cryptomys may have been a result of environmental changes associated with rifting.

Morphological castes in a vertebrate.

Morphological specialization for a specific role has, until now, been assumed to be restricted to social invertebrates. Herein we show that complete physical dimorphism has evolved between reproductives and helpers in the eusocial naked mole-rat. Dimorphism is a consequence of the lumbar vertebrae lengthening after the onset of reproduction in females. This is the only known example of morphological castes in a vertebrate and is distinct from continuous size variation between breeders and helpers in other species of cooperatively breeding vertebrates. The evolution of castes in a mammal and insects represents a striking example of convergent evolution for enhanced fecundity in societies characterized by high reproductive skew. Similarities in the selective environment between naked mole-rats and eusocial insect species highlight the selective conditions under which queen/worker castes are predicted to evolve in animal societies.

Circulating LH levels and the response to exogenous GnRH in the common mole-rat: implications for reproductive regulation in this social, seasonal breeding species.

The effects of breeding season and reproductive status on male and female reproduction were investigated in the common mole-rat, Cryptomys hottentotus hottentotus, a cooperatively breeding rodent which exhibits a unique combination of seasonal breeding and a reproductive division of labor. Pituitary function was examined by measuring the luteinizing hormone (LH) responses to single doses of 2 microg exogenous gonadotrophin-releasing hormone (GnRH) and physiological saline in 69 males and 58 females from 35 wild caught colonies. Neither males nor females exhibited any apparent manifestation of season on basal LH concentrations or on pituitary sensitivity to stimulation by exogenous GnRH. The continuance of reproductive function during the nonbreeding period is essential in common mole-rat males and females, as this period coincides with the period of maximal dispersal opportunity in the winter rainfall area they inhabit. Normal circulating levels of reproductive hormones in dispersing animals may aid intersexual recognition, assist pairbond formation, and thus prime animals for independent reproduction. Circulating basal concentrations of LH as well as LH levels measured in response to a single exogenous GnRH challenge were not significantly different between the reproductive and non-reproductive groups of either sex, suggest the absence of a physiologically well-defined suppression of reproduction in subordinate common mole-rats.

Ecological constraints drive social evolution in the African mole-rats.

The African mole-rats (family Bathyergidae) are subterranean hystricomorph rodents occurring in a variety of habitats and displaying levels of sociality which range from solitary to eusocial, making them a unique mammalian taxonomic group to test ecological influences on sociality. Here, we use an extensive DNA-based phylogeny and comparative analysis to investigate the relationship between ecology, sociality and evolution within the family. Mitochondrial cytochrome-b and 12s rRNA trees reveal that the solitary species are monophyletic when compared to the social species. The naked mole-rat (Heterocephalus glaber) is ancestral and divergent from the Damaraland mole-rat (Cryptomys damarensis), supporting previous findings that have suggested the multiple evolution of eusociality within the family. The Cryptomys genus is species-rich and contains taxa exhibiting different levels of sociality, which can be divided into two distinct clades. A total of seven independent comparisons were generated within the phylogeny, and three ecological variables were significantly correlated with social group size: geophyte density (p < 0.05), mean months per year of rainfall greater than 25 mm (p < 0.001), and the coefficient of rainfall variation (p = 0.001). These results support the food-aridity hypothesis for the evolution of highly social cooperative behaviour in the Bathyergidae, and are consistent with the current theoretical framework for skew theory.

A dispersive morph in the naked mole-rat.

Close inbreeding is known for a variety of small mammal species for which a high probability of mortality during dispersal makes helping and delayed maturation a relatively secure fitness option. Prolonged inbreeding, however, is usually associated with lowered fitness, and it has been shown that most highly inbred small mammals and social insects have inbreeding-avoidance mechanisms that promote some degree of outbreeding. However, previous field and laboratory research on the naked mole-rat (Heterocephalus glaber) suggested that this cooperatively breeding rodent is highly inbred, with new colonies forming by fission. Here we report the discovery of a dispersal phenotype that may occasionally promote outbreeding in naked mole-rats. These dispersers are morphologically, physiologically and behaviourally distinct from other colony members. They are laden with fat, exhibit elevated levels of luteinizing hormone, have a strong urge to disperse, and only solicit matings with non-colony members. These findings suggest that, although rare, a dispersive morph exists within naked mole-rat colonies.

Subterranean mole-rats naturally have an impoverished calciol status, yet synthesize calciol metabolites and calbindins.

Mole-rats (Family Bathyergidae) have no obvious source of calciol. They live in an environment devoid of sunlight and consume a herbivorous diet. Calciol status, metabolism and expression were examined in six species of Bathyergids. Serum levels of calcidiol in all species were < 5 micrograms/l and those of calcitriol were low (18.0 +/- 11.0 (SD) ng/l, N = 57) when compared to other rodents. Within 72 h of injecting animals with tritium-labelled calciol, most of the labelled prohormone had been metabolized to more polar metabolites. Three times more tritium-labelled calcitriol (19.3 +/- 2.9%) was present than (24R)-hydroxycalcidiol (6.2 +/- 10%). The natural absence of detectable circulating concentrations of calcidiol and the threefold greater amount of calcitriol to (24R)-hydroxycalcidiol produced indicate that calciol naturally is in short supply. Calciol-dependent calbindins were absent in the duodenum. Calbindin-D28k was present in the Purkinje cells of the cerebellum and in some collecting ducts and proximal and distal convoluted tubules of the kidney. Calbindin-D9k also was present but was localized uniquely in the juxtaglomerular cells of the five southern African species. These data confirm that Bathyergid mole-rats naturally have an impoverished calciol status. Despite the presence of calbindins in renal tissues, the functional importance of this hormone in calbindin synthesis and other normal mole-rat physiology is not known.

Investigation of numbers and motility of spermatozoa in reproductively active and socially suppressed males of two eusocial African mole-rats, the naked mole-rat (Heterocephalus glaber) and the Damaraland mole-rat (Cryptomys damarensis).

Reproductive tracts and spermatozoa from reproductively active and reproductively suppressed non-breeding males from two species of eusocial African mole-rats Cryptomys damarensis and Heterocephalus glaber were examined. In two captive colonies of Heterocephalus glaber, reproductive tracts from seven non-breeding males removed from their colonies, and housed singly for 5-6 weeks to cause reproductive activation, were compared with reproductive tracts from seven non-breeding males. The body weight of the separated, reproductively active males increased significantly (P < 0.01), and the mean testis weights relative to body weight of the reproductively active males were significantly larger (P < 0.05) than those of non-breeding males. The number of spermatozoa, in one half of the reproductive tract, was higher in active males than in non-breeding males (mean +/- SEM: 8.59 x 10(6) +/- 2.69 x 10(6) versus 1.78 x 10(6) +/- 1.43 x 10(6), respectively; P < 0.05). In addition, six of the seven reproductively active males, but only two of seven non-breeding males, had motile spermatozoa. A total of 28 wild Cryptomys damarensis from two colonies were examined in the field. The testis weights relative to body weight of breeding males (n = 7) were higher than those of non-breeding males (n = 19; P < 0.01), but the number of spermatozoa did not differ significantly between the two groups (0.13 x 10(6) +/- 0.06 x 10(6), n = 7 versus 0.29 x 10(6) +/- 0.14 x 10(6), n = 21, respectively). Breeding and non-breeding males produced similar numbers of motile spermatozoa.(ABSTRACT TRUNCATED AT 250 WORDS)

Thermoregulation in three species of Afrotropical subterranean mole-rats (Rodentia: Bathyergidae) from Zambia and Angola and scaling within the genus Cryptomys.

The thermoregulatory characteristics of three species of Cryptomys from Zambia and Angola are examined and, together with published data on four other species of Cryptomys from southern Africa, used to determine whether scaling occurs in this genus of subterranean rodents. The thermoregulatory properties of acclimated giant Zambian mole-rats, Cryptomys mechowi ([Formula: see text]=267 g), Angolan mole-rats, Cryptomys bocagei ([Formula: see text]=94 g) and Zambian common mole-rats Cryptomys hottentotus amatus ([Formula: see text]=77 g) are as follows. Mean resting metabolic rates (RMRs) within the respective thermoneutral zones were 0.60±0.08 cm3 O2 g-1 h-1 (n=12) for C. mechowi; 0.74±0.06 cm3 O2 g-1 h-1 (n=8) for C. bocagei and 0.63±0.06 cm3O2 g-1 h-1 (n=21) for C. h. amatus. The thermoneutral zones (TNZs) of all three species are narrow: 29-30°C for C. mechowi; 31.5-32.5°C for C. bocagei and 28-32° C for C. h. amatus. The increase in mean RMR at the lowest temperatures tested (15° C for C. mechowi, 18° C for C. bocagei and C. h. amatus) was 2.35, 2.2 and 3.82 times their RMR in the TNZ respectively. Body temperatures are low, 34±0.53° C (n=24) for C. mechowi, 33.7±0.32° C (n=20) for C. bocagei and 33.8±0.43° C (n=40) for C. h amatus. At the lower limit of thermoneutrality, conductances are 0.09±0.01 cm3 O2 g-1 h-1 °C-1 (n=30) in C. mechowi; 0.12±0.01 cm3 O2 g-1 h-1 °C-1 (n=20) in C. bocagei and 0.12±0.03 cm3 O2 g-1 h-1 °C-1 (n=32) in C. h. amatus. The range in mean body mass among the seven species of Cryptomys examined for scaling was 60 g (C. darlingi) to 267 g (C. mechowi). There is no clear relationship between RMR within the TNZ and body mass. The resultant relationship is represented by the power curve RMR=2.45 mass-0.259.

Mammalian eusociality: a family affair.

Comparative studies of two species of mole-rat are helping to clarify the ecological correlates of mammalian eusociality. Both species live in social groups composed of close kin, within which breeding is restricted to one female and one to three males. They inhabit xeric areas with dispersed, patchy food and unpredictable rainfall. During droughts, they can neither expand their tunnel systems nor disperse. In brief periods after rain the animals must cooperate and dig furiously to locate rich food patches. By living in groups, arid-zone mole-rats can take full advantage of windows of opportunity when conditions are right for burrowing. Thus, ecological factors and kin selection have apparently interacted in the evolution of eusociality in these species.

LH responses to single doses of exogenous GnRH by freshly captured Damaraland mole-rats, Cryptomys damarensis.

Pituitary function in reproductive and nonreproductive colony members of Damaraland mole-rats, Cryptomys damarensis, was investigated by measuring the LH responses to single doses of 2 micrograms exogenous GnRH and physiological saline in 29 females and 37 males (31 of these animals from two entire colonies). In females, basal LH concentrations were significantly greater in reproductive (n = 9) than in nonreproductive animals (n = 11): 7.6 +/- 1.0 versus 4.3 +/- 0.6 miu ml-1, respectively (P < 0.001). Reproductive females had a significantly greater LH response to 2.0 micrograms GnRH (7.6 +/- 1.0 to 37.7 +/- 6.2 miu ml-1; n = 9) than did nonreproductive females (4.3 +/- 0.6 to 11.8 +/- 1.0 miu ml-1; n = 11, P < 0.001). In contrast, there was no significant difference in basal LH concentrations between reproductive (n = 8) and nonreproductive males (n = 20): 5.3 +/- 4.3 versus 3.2 +/- 1.2 miu ml-1, respectively. There was also no difference in LH response to the administration of 2.0 micrograms GnRH between reproductive and nonreproductive males (5.3 +/- 4.3 to 21.8 +/- 8.6 miu ml-1; n = 8; versus 3.2 +/- 1.2 to 21.1 +/- 8.5 miu ml-1; n = 21; P = 0.5). When the results from the two entire colonies were analysed separately, LH responses to GnRH in the 11 nonreproductive females were less than in the two reproductive females. In contrast, the response of two reproductive males in the colonies did not differ from that of 16 nonreproductive males, although these latter comparisons could not be validated statistically.(ABSTRACT TRUNCATED AT 250 WORDS)

Seismic signal transmission between burrows of the Cape mole-rat, Georychus capensis.

Both seismic and auditory signals were tested for their propagation characteristics in a field study of the Cape mole-rat (Georychus capensis), a subterranean rodent in the family Bathyergidae. This solitary animal is entirely fossorial and apparently communicates with its conspecifics by alternately drumming its hind legs on the burrow floor. Signal production in this species is sexually dimorphic, and mate attraction is likely mediated primarily by seismic signalling between individuals in neighboring burrows. Measurements within, and at various distances away from, natural burrows suggest that seismic signals propagate at least an order of magnitude better than auditory signals. Moreover, using a mechanical thumper which could be triggered from a tape recording of the mole-rat's seismic signals, we established that the vertically-polarized surface wave (Rayleigh wave) propagates with less attenuation than either of the two horizontally-polarized waves. Thus, we tentatively hypothesize that Rayleigh waves subserve intraspecific communication in this species.

Social suppression of reproduction in male naked mole-rats, Heterocephalus glaber.

To investigate possible anatomical and endocrine differences between breeding and non-breeding male naked mole-rats, 113 animals from 24 captive and 4 wild colonies were studied. While breeding males had larger reproductive tract masses compared to non-breeders relative to body mass (P less than 0.01), spermatogenesis was active in all of the non-breeding males examined histologically (n = 9) and spermatozoa were present in the epididymides. Compared with non-breeders, breeding males had significantly higher urinary testosterone concentrations (mean +/- s.e.m.: 23.8 +/- 2.3 vs 5.2 +/- 1.4 ng/mg Cr respectively; P less than 0.001), and plasma LH (10.7 +/- 1.7 vs 5.0 +/- 0.8 mi.u./ml respectively; P less than 0.01). Single doses of 0.1, 0.5 or 1.0 microgram GnRH produced a significant rise in plasma LH concentrations 20 min after s.c. injection in breeding and non-breeding males at all doses (P less than 0.001). However, there were differences in the magnitude of the LH response following administration of GnRH between breeding and non-breeding males, with non-breeding males showing a dose-response and having lower plasma LH concentrations 20 min after a single injection of 0.1 or 0.5 microgram (P less than 0.05), but not 1.0 microgram, GnRH. This apparent lack of pituitary sensitivity of non-breeding males to single doses of exogenous GnRH was reversed by 4 consecutive injections of 0.5 microgram GnRH at hourly intervals, suggesting that the reduced sensitivity may be the result of insufficient priming of the pituitary by endogenous GnRH. These results indicate that, despite the fact that non-breeding males were apparently producing mature gametes, clear endocrine deficiencies existed in male naked mole-rats.

LH responses of female naked mole-rats, Heterocephalus glaber, to single and multiple doses of exogenous GnRH.

To investigate possible differential pituitary secretion of LH in breeding and non-breeding female naked mole-rats, the LH responses to administration of exogenous GnRH were measured in 55 females from 20 captive colonies. Single doses of 0.1, 0.5 or 1.0 micrograms GnRH produced a significant rise in plasma LH concentrations 20 min after s.c. injection in breeding and non-breeding females at all doses (P less than 0.001). While at the highest dose of 1.0 microgram there was no difference in the LH response between breeding and non-breeding females, as the dose was lowered there was a progressive decline in the LH response in non-breeding females such that, at the 0.1 microgram dose, GnRH produced only a small, but significant, increase in plasma LH (1.3 +/- 0.2 to 2.9 +/- 0.5 mi.u./ml, N = 5) compared with breeding females (3.4 +/- 0.8 to 9.6 +/- 2.0 mi.u./ml, N = 6). The LH responses of the latter were not significantly reduced at the lower doses of GnRH. The apparent lack of sensitivity to low doses of exogenous GnRH in non-breeding females was reversed by 4 consecutive 1-h injections of 0.1 microgram, which produced a rise in LH from 1.2 +/- 0.2 to 9.0 +/- 0.2 mi.u./ml (N = 4), comparable to that of breeding females given a single injection of 0.1 microgram GnRH. These results suggest that the anterior pituitary in non-breeding female naked mole-rats is less sensitive to low doses of exogenous GnRH than in breeding females, possibly due to a lack of priming by endogenous GnRH. Therefore, the socially-induced block to ovulation in non-breeding female naked mole-rats may be due to inhibition of hypothalamic GnRH secretion.

Social suppression of ovarian cyclicity in captive and wild colonies of naked mole-rats, Heterocephalus glaber.

To investigate the endocrine cause of reproductive suppression in nonbreeding female naked mole-rats, animals from 35 colonies were studied in captivity. Urinary and plasma progesterone concentrations were elevated in pregnant females (urine: 10.0-148.4 ng/mg Cr, 27 samples from 8 females; plasma: 3.6-30.0 ng/ml, 5 samples from 5 females; Days 21-40 of pregnancy) and cyclic breeding females (urine: 0.5-97.8 ng/mg Cr, 146 samples from 7 females; plasma: less than 1.0-35.4 ng/ml, 25 samples from 7 females). The latter group showed cyclic patterns of urinary progesterone, indicating a mean ovarian cycle length of 34.4 +/- 1.6 days (mean +/- s.e.m.) with a follicular phase of 6.0 +/- 0.6 days and a luteal phase of 27.5 +/- 1.3 days (19 cycles from 9 breeding females). In non-breeding females urinary and plasma progesterone values were undetectable (urine: less than 0.5 ng/mg Cr, 232 samples from 64 females; plasma: less than 1.0 ng/ml, 7 samples from 6 females). Breeding females had higher (P less than 0.001) plasma LH concentrations (3.0 +/- 0.2 mi.u./ml, 73 samples from 24 females) than did non-breeding females (1.6 +/- 0.1 mi.u./ml, 57 samples from 44 females). Urinary and plasma progesterone concentrations in non-breeding females from wild colonies situated near Mtito Andei, Kenya, were either below the assay sensitivity limit (urine: less than 0.5 ng/mg Cr, 11 females from 2 colonies; plasma: less than 1.0 ng/ml, 25 females from 4 colonies), or very low (plasma: 1.6 +/- 0.6 ng/ml, 15 females from 4 colonies). In captivity, non-breeding females removed from their colonies (i.e. the dominant breeding female) and either paired directly with a non-breeding male (N = 2), or removed and housed singly for 6 weeks before pairing with a non-breeding male (N = 5) may develop a perforate vagina for the first time in as little as 7 days. Urinary progesterone concentrations rose above 2.0 ng/mg Cr (indicative of a luteal phase) for the first time 8.0 +/- 1.9 days after being separated. These results suggest that ovulation is suppressed in subordinate non-breeding female naked mole-rats in captive and wild colonies, and show that plasma LH concentrations are significantly lower in these non-breeding females. This reproductive block in non-breeding females is readily reversible if the social factors suppressing reproduction are removed.

Identification of crystalline allantoin in the urine of African Cricetidae (Rodentia) and its role in their water economy.

All eleven cricetid species, examined in this investigation, produced an off-white crystalline precipitate in their urine when deprived of water, whereas not one murid examined did so. This crystalline compound was identified as allantoin, a common end product of purine catabolism. The quantity found in the solid precipitate alone accounted for 47% of the total nitrogen excreted and was approximately 14 times greater than the predicted quantity of allantoin from purine degradation. It appears that there is a shift in nitrogen excretion from urea to allantoin in the Cricetidae. Water-deprived cricetids had higher urine osmolalities, urea concentrations and lower daily percentage body water turnovers than the murids. This can be explained by the substantial water savings associated with excreting solid allantoin. The discrepancy in the mode of nitrogen excretion between the two families inhabiting the Namib Desert can be attributed to their different evolutionary histories, the Cricetidae being pre-adapted for survival in deserts.

Nutrition and burrowing energetics of the Cape mole-rat Georychus capensis.

At 22°C the resting oxygen consumption of G. capensis is 1.13±0.05 cm3O2·g-1·h-1 (mean± S.E.). In loose sandy soil the burrowing metabolic rate was approximately three times that of resting (3.41±0.19 cm3O2·g-1· h-1). Rate of oxygen consumption while burrowing bears a linear relationship with rate of burrowing. The equation of the regression line describing this relationship was used to construct a model for calculating energy expenditure of burrowing in free-living mole-rats. The diet of G. capensis consists of some green plant material and geophyte corms. The latter has a mean gross energy content of 16.36 kJ·g-1 dry weight. The digestibility coefficient for captive G. capensis fed on sweet potato, was 97.42±0.41%. Data collected from an excavated burrow system revealed that the total energetic cost of constructing the burrow amounted to 79% of the estimated digestible energy available from geophyte corms in the area. A food store in the same burrow system was sufficient to meet the maintenance requirements of an adult G. capensis, resting at 22°C, for approximately 80-85 days. Soil samples taken at random adjacent to the burrow contained corms with a mean estimated digestible energy value of 2084 kJ per m3 of soil. A comparison of energetic cost of burrowing and randomly available digestible energy in the field suggests that foraging patterns are not random.

Eusociality in a mammal: cooperative breeding in naked mole-rat colonies.

Laboratory observations on a field-collected colony of 40 Heterocephalus have shown that only a single female breeds. The remaining individuals constitute two or three castes, each containing both sexes and distinguishable by size differences and the tasks they perform. These features, together with long life-spans, overlap of generations, cooperative brood care, and possible age polyethism provide parallels with the eusocial insects.