The retina consists of many parallel circuits designed to maximize the gathering of important information from the environment. Each of these circuits is comprised of a number of different cell types combined in modules that tile the retina. To a subterranean animal, vision is of relatively less importance. Knowledge of how circuits and their elements are altered in response to the subterranean environment is useful both in understanding processes of regressive evolution and in retinal processing itself. We examined common cell types in the retina of the naked mole-rat, Heterocephalus glaber with immunocytochemical markers and retrograde staining of ganglion cells from optic nerve injections. The stains used show that the naked mole-rat eye has retained multiple ganglion cell types, 1–2 types of horizontal cell, rod bipolar and multiple types of cone bipolar cells, and several types of common amacrine cells. However, no labeling was found with antibodies to the dopamine-synthesizing enzyme, tyrosine hydroxylase. Although most of the well-characterized mammalian cell types are present in the regressive mole-rat eye, their structural organization is considerably less regular than in more sighted mammals. We found less precision of depth of stratification in the inner plexiform layer and also less precision in their lateral coverage of the retina. The results suggest that image formation is not very important in these animals, but that circuits beyond those required for circadian entrainment remain in place.

The Namaqua dune mole-rat Bathyergus janetta breeds seasonally, producing up to two litters during the early summer in the Cape. Females exhibit elevated urinary progesterone and oestradiol-17β concentrations and males raised concentrations of testosterone, these correlate with the onset of the seasonal winter rainfall. The rains moisten the soil allowing excavation and provide the mole-rats with the opportunity not only to extend burrow systems but also to search for mates. An increased incidence of hindfoot drumming and visitations to neighbouring burrow systems occurs during this period. Plural occupancy of burrows by adult mole-rats was found during the onset of the winter, and pups, lactating and pregnant females were caught at the end of October and November (early Cape summer). Pups develop rapidly and their eyes open after c. 1 week. Pup body mass increases exponentially from birth to about 70 days. Inter-sibling aggression increases progressively and after c. 60 days captive-born pups had to be separated, suggesting that in the wild they would have dispersed.

Subterranean mammals inhabit an environment that is normally devoid of light and are therefore deprived of photoperiodic information that can be used to time important life-history events. An assessment was made of whether melatonin secretion in a strictly subterranean rodent, the Damaraland mole-rat Cryptomys damarensis, can be modified by photoperiod. In experiment 1, a clear diurnal rhythm of melatonin secretion in animals housed under a neutral photoperiod (12L:12D) was observed, with significantly higher melatonin concentrations in the dark compared to the light phase. The same diurnal melatonin rhythm was found 1 day after animals were transferred to either continuous light or continuous dark, suggesting that a circadian rhythm was maintained under acute exposure to light and dark. In experiment 2, melatonin secretion was monitored in a long (14L:10D) and short day (10L:14D) photoperiod and was found to be modified by the photoperiodic change. We therefore suggest that the Damaraland mole-rat possesses a circadian melatonin rhythm that can be physiologically modulated in response to photoperiod.

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 highveld mole-rat, Cryptomys hottentous pretoriae is a social, subterranean, co-operative breeder with a high reproductive skew. The relationship between reproductive status and reproductive physiology in male highveld mole-rats was investigated. Testicular morphometrics, histological parameters, plasma testosterone concentrations and sperm motility parameters were studied in 31 males (14 reproductive and 17 non-reproductive males). Reproductive males were significantly larger than non-reproductive males, with testicular mass and volume corrected for body mass considerably larger for the reproductive males. Circulating plasma testosterone concentrations of reproductive males were not significantly higher than non-reproductive males (reproductive males 10.3±1.8 nmol l−1vs non-reproductive males 7.3±1.74 nmol l−1). Sperm motility parameters were measured, but no significant differences were found between reproductive and non-reproductive males. All 14 reproductive males had motile sperm, whereas 13 of the 17 (76%) of non-reproductive males possessed motile sperm. A typical ejaculate of a reproductive male contains 48.3% motile sperm characterized by a high percentage of flagellar defects, whereas the non-reproductive male has an ejaculate containing 45.6% motile sperm with a high percentage of head defects. It is apparent that non-reproductive males are not physiologically suppressed from reproducing. Moreover, the non-reproductive males are excluded from incestuous matings as a result of being the offspring of the reproductive female.

Some aspects of the colony and social structure of the Ghana mole-rat Cryptomys zechi were studied in a mesic Guinea savanna woodland in Ghana. Colony size ranged between one and seven (average 3.6). The mean adult body weight was 217.9 g and the colony biomass was 634 g. Responsibility for colony safety in C. zechi, as revealed from order of capture in response to damage to the burrow system, is age dependent, with sub-adults being the most frequent repairers. There are indications that male workers are more involved in colony maintenance than females and that the reproductive individuals, especially the males, perform little work. We discuss the finding that C. zechi although a social bathyergid, is close to the bottom of the sociality ladder.

Blind mole-rats (Spalax ehrenbergi) are fossorial solitary rodents that present striking morphological, physiological and behavioural adaptations to the subterranean environment in which they live. Previous studies have shown that mole-rats are specialised in tooth-digging. The rapid eruption-rate of their incisors has evolved to compensate for their excessive wear by excavation. Males use their incisors more than females for digging and fighting, and their rate of incisor eruption is significantly more rapid than in females. Since mole-rats use their incisors for digging throughout the year, we suggest that continuous mechanical pressure on their oral tissues concentrated at the apical sites of the upper incisors leads to cell and tissue fatigue. We provide evidence for 5 stages of palatal perforation by the upper incisors at their apical sites, with maximum perforation characterising aged males. Interspecies comparisons with 7 other fossorial and semi-fossorial rodent species, and with beavers, which expose their incisors to enormous mechanical pressure, revealed that this palatal perforation is unique to the male mole-rat. We suggest that while the fast eruption rate of incisors in the mole-rat compensates for the rapid wear resulting from digging, evolutionary adaptation to continuous tooth-digging is still ongoing, since the physical pressure of digging at the apical sites of the upper incisors leads to tissue destruction, breakage of the palatal bone and possibly to death, as a result of maxillary inflammation.

Colonies of the Mashona mole-rat Cryptomys darlingi are founded from a single reproductive pair of animals that are genetically unrelated by descent. All non-reproductive animals are the progeny of the reproductive pair. Non-reproductive colony members do not seem to be suppressed from reproduction at the level of the pituitary. In colonies in which the reproductive female is removed or dies, there is strict incest avoidance and the colony remains reproductively quiescent. Reinstatement of sexual activity in a queenless colony may be brought about in the laboratory by the introduction of unfamiliar and unrelated adult males. In the queenless colony under study, there was a marked change in the dominance hierarchy with an increase in Landau's index of linearity from 0.61 to 1.0 on the introduction of two unrelated males. The two new males became the most dominant within the colony. All previously non-reproductive females exhibited heightened urinary oestradiol 17β and progesterone concentrations on the introduction of the males. However, it was only the older and most dominant non-reproductive female that became the new reproductive female and produced a litter of three pups 70 days after the initial introduction.

The highveld mole-rat Cryptomys hottentotus pretoriae exhibits an extreme reproductive division of labour. Reproduction in this seasonally breeding social mole-rat is restricted to a breeding pair. The non-reproductive females seem to be physiologically suppressed from reproducing, whereas the non-reproductive males are restricted from sexual activity through incest avoidance. Circulating basal concentrations of luteinizing hormone (LH) between reproductive and non-reproductive females were not significantly different, yet the response to an exogenous gonadotrophin-releasing hormone (GnRH) challenge was significantly lower in non-reproductive females when compared to reproductive females. The mean basal LH concentration in non-reproductive females was not significantly different in and out of the breeding season. However, the magnitude of the LH response in non-reproductive females out of the breeding season was greater than in the breeding season. The ovaries of reproductive females showed little change with respect to season. However, reproductive females had enhanced follicular activity in comparison to their non-reproductive counterparts. This suggests that non-reproductive females are under stricter control from the reproductive female when she is pregnant. The tropical African mole-rats exhibit a continuum of socially induced infertility with differing social species inhabiting regions of varying degrees of aridity. In this spectrum, a predominantly behavioural repression is found amongst non-reproductive females in social mesic-adapted species, whereas a strictly physiological suppression is operative amongst non-reproductive females in arid-adapted social species. Here we show that the mesic-adapted highveld mole-rat does not fit this trend in that the non-reproductive females seem to be physiologically suppressed. We discuss this in light of the seasonality of breeding and the ecological constraints that may operate during times of drought.

Temperature and respiratory gas (CO2 and O2) concentrations were measured in the foraging tunnels of burrows naturally inhabited by two species of southern African mole-rats, the Cape mole-rat Georhychus capensis and the Damaraland mole-rat Cryptomys damarensis. Both species are completely fossorial and inhabit closed burrow systems. Tunnels of G. capensis burrows had a mean diameter of 8.7 cm and a depth, measured to the roof of the tunnel, of 6.2 cm; those of C. damarensis had a mean diameter of 6.5 cm and depth of 40 cm. In both species, the mean concentration of CO2 was higher, and mean concentration of O2 lower, in burrows than in the surrounding soil or in ambient air. Mean and minimum values of O2 were 20.4% and 19.8%, respectively, in G. capensis and 20.4% and 19.9% in C. damarensis; mean and maximum values of CO2 were 0.4% and 1.2% in G. capensis and 0.4% and 6.0% in C. damarensis. Temperature varied between 18.5 and 24.2 °C in burrows of G. capensis by comparison with an ambient range of 16.9 to 26.8 °C; and from 19.6 to 29.3 °C in burrows of C. damarensis by comparison with an ambient range of 8.6 to 30.8 °C. Thus a burrowing habit seems to offer both species protection from extremes of temperature without entailing the cost of a grossly abnormal respiratory environment. From a review of the relevant literature, we conclude that average concentrations of CO2 and O2 in mammalian burrows often do not differ greatly from ambient values. However, more work is needed to determine the respiratory gas concentrations in the immediate vicinity of active, burrowing animals.

The giant mole-rat Cryptomys mechowi exhibits an extreme reproductive division of labour. Reproduction in the colony is restricted to a single breeding pair. Non-reproductive individuals do not breed unless removed from the colony and paired with unfamiliar conspecifics. Circulating basal concentrations of luteinizing hormone (LH) as well as LH levels measured in response to a single exogenous gonadotropin hormone (GnRH) challenge are not significantly different between the reproductive and non-reproductive groups of either sex. Neither non-reproductive males nor females are physiologically suppressed at the level of the pituitary, as they do not exhibit a reduced pituitary secretion of LH or decreased sensitivity to hypothalamic GnRH. Rather, reproductive skew seems to result from an inhibition to mate with familiar conspecifics. Familiar individuals housed in the absence of the breeding pair did not breed, whereas pairing unfamiliar individuals of the opposite sex resulted in sexual activity and recruitment. The results from the giant mole-rat suggest that reproductive skew may result from a predominantly behavioural repression in non-reproductive females arising from an incest avoidance mechanism.

A total of 104 Damaraland mole-rats was collected in 1988, 1990, 1991 and 1993 from an open water canal in the Otjiwarongo district of Namibia. The mole-rats were apparently dispersing above ground when they fell in and drowned; they had travelled at least 20 m in distance from their natal burrow systems. The dispersing mole-rats of both sexes were significantly larger than 73 animals trapped from a nearby control population. Sequential tooth wear and eruption patterns showed that dispersers were adult animals. There was a bias towards male mole-rats dispersing. Furthermore, in years with lower rainfall (< 300 mm) it was found that dispersal became even more male biased. Eighty per cent of males and 40% of female dispersers showed signs of reproductive readiness, with more males than females being ready to breed. Thus it appears that, in these otherwise totally subterranean mammals, at least one form of dispersal occurs, above ground, and that adult mole-rats are the principal dispersers.

The relationship between reproductive status and reproductive physiology was investigated in male Damaraland mole-rats Cryptomys damarensis, an eusocial rodent that exhibits a marked reproductive division of labour. Testicular anatomical and histological morphometrics, selected sperm motion parameters and measurements of circulating plasma testosterone were studied in 44 males (17 reproductive and 27 non-reproductive). Reproductive males were significantly larger than non-reproductive males. Testicular mass of reproductive males, corrected for body mass, was significantly larger than in non-reproductives. However, corrected testicular volume did not vary significantly between the two groups. Circulating plasma testosterone concentrations of reproductives and non-reproductives were not significantly different (reproductive males 201.5 ± 56.5 pg/l vs 187.5 + 91 pg/l for non-reproductive males). The quality of sperm was measured by the percentage of spermatozoa produced and the amount of immature sperm, and differed between the two groups. Reproductive males produced a greater proportion of motile sperm (77.8%) than non-reproductives (56.3%). Non-reproductive males were characterized by a greater proportion of immature sperm in the epididymis (79%) than the reproductives (25%). Interestingly, the spermatozoa of reproductive males were characterized by a greater percentage of head and tail defects. Oligospermia may be the result of a reduction in circulating follicle stimulating hormone, which acts upon the Sertoli cells of the seminiferous tubules, that in turn may be due to a lack of an opportunity to mate while in the confines of the colony in this obligate outbreeder.

The highveld mole-rat Cryptomys hottentotus pretoriae occurs in the summer rainfall region of the highveld of South Africa. It lives in colonies of up to 10 individuals, in which reproduction is limited to one to two of the largest males and the largest female in the colony. Of three field captured colonies, the mean colony size was 10 ± 0 animals with a mean biomass of 900 ± 147 g. The study colony had a sex ratio of 1.5:1 in favour of males.

The social hierarchy of the highveld mole-rat is non-linear and has a value of 0.3 calculated from Landau's linearity index. Dominance was negatively correlated with capture order.

Histological examination of the gonads of reproductively active and sexually quiescent members of both sexes of two colonies provides further support of there being a single reproductive male and female responsible for procreation in the colony.

The non-reproductives of both sexes appear to be important in burrow maintenance activity and in foraging for geophytes upon which the mole-rats feed. The non-reproductive members of the colony cannot be subdivided into different worker groups based on body mass. The available evidence suggests that the highveld mole-rat is a transiently social species, at one extreme of a sociality continuum in the genus Cryptomys. The position of this subspecies in respect to the sociality continuum is discussed.

In this study factors that influence the plasticity in growth and adult body mass in the naked mole-rat (Heterocephalus glaber) are explored. Growth functions (Gompertz transformation) revealed that naked mole-rat pups have the slowest mean maximum growth rate (0.207 g.day−1) of all the Bathyergidae. Growth functions varied significantly between litters and there was an inverse trend between asymptotic body mass and litter order. Similarly, the magnitude of the growth response following the death of the breeding male or older siblings was greater for older litter members. There was a significant correlation between body mass and age at all stages of the colony's history although this relationship weakened with time. It is suggested that dominance interactions between old and young mole-rats within the colony are important factors influencing the growth and reproductive potential of all colony members. Changes in body mass were linked to the behaviour of the individual within the colony providing a mechanism for the apparent match between large body size and colony defence and small body size and work frequency. Despite low genetic variation within colonies there is considerable variation in the size of colony members suggesting that plasticity in growth is the product of different ontogenetic histories of colony members that may serve as the basis for variation in size amongst adults.

Crytomys damarensis is a eusocial subterranean rodent occurring in colonies in which a sole reproductive female and one to two males are involved in procreation; the remaining colony members are non-reproductive. Within each sex the reproductive animals are usually the largest and most dominant animals.

The dominance hierarchy of a young colony consisting of a founding reproductive pair of genetically unrelated animals and the first three successive litters, comprising 12 offspring, from this pair had a non-linear dominance hierarchy, h = 0.32. Dominance was found to be correlated positively to body mass as well as to the age of the animal. A subsequent study, which followed the incorporation of two subsequent litters and growth of the first three litters, and comprised 15 offspring, revealed that the dominance hierarchy had attained linearity, h = 0.9. Dominance was again positively correlated with the age and body mass of respective colony members. In both studies there was no clear relationship between dominance and sex within the colony.

The relationship between body mass and burrow-maintenance activity revealed that frequent workers in the colony had a greater mean body mass than infrequent workers. It is suggested that in establishing colonies, the older, stronger and hence larger animals may constitute the initial workforce. In addition the colony exhibits frequency-related work determination into frequent and infrequent workers. Gender was found to play no role in the frequency of work associated with burrow maintenance.

The social organization and temporal changes with recruitment are discussed in the light of events occurring in the field.

Cryptomys mechowi is a social subterranean mole-rat which inhabits the mesic regions of south-eastern and central Africa. Giant mole-rats live in colonies of up to 11 animals in which reproduction is normally restricted to the largest male and female in the colony. In this preliminary study on a laboratory-reared colony, the non-reproductive members of the colony could not be placed into clearly defined, work-related groups based on body mass. The colony showed a work frequency related division of labour. Reproductive animals performed little work, 3-4% of the total work. The non-reproductive caste was divided into two groups based on the amount of burrow maintenance activity: those performing between 7-11% of total work and those ranging from 15-22% of total work. Male members of the colony tended to perform more work than the females. The dominance hierarchy of the colony was found to be nearly linear, with a value of 0.83 calculated from Landau's linearity index. The reproductive animals were the most dominant in their respective gender. Dominance was not age-related. However, non-reproductive males tended to be more dominant than non-reproductive females. Social organization within the giant Zambian mole-rat is compared with other southern African Cryptomys species in the light of the eusociality continuum.

Cryptomys hottentotus has no access to dietary or endogenous sources of cholecalciferol (D3). Serum concentrations of calcifediol (25(OH)D3) were undetectable (< 5 ng/ml) and calcitriol (1,25(OH)2D3), although detectable in plasma, was at a low concentration (31.40) (SEM 3.06 pg/ml). Despite their naturally impoverished vitamin D status, these animals exhibited highly efficient mineral absorption processes (Ca apparent fractional absorption efficiency, AFA (%) 95.33 (SEM 1.36); inorganic P (Pi) AFA (%) 93.49 (SEM 0.81)). Furthermore, plasma mineral content was tightly regulated (Ca 2.57 (SEM 0.08) mmol/l; Mg 1.23 (SEM 0.05) mmol/l; Pi 2.12 (SEM 0.15) mmol/1). Mode of uptake in C. hottentotus was unlike that in other D3-replete mammals. First, passive (rather than active) uptake occurred in the traditional site of active Ca absorption (with serosal: mucosal (S:M) ratios in the duodenum of 1.32 (SEM 0.13)), and the only site of active Ca uptake was the hindgut (caecum S:M 3.35 (SEM 0.46); proximal colon S:M 2.13 (SEM 0.30)). Despite the presence of active uptake in these hindgut regions, their overall contribution to the daily rate of mineral absorption was small (9.53 (SEM 1.27)%). These underground inhabitants rely upon highly efficient, passive mineral uptake. This is adequate to meet their mineral requirements and maintain mineral homeostasis in the absence of vitamin D.

The effect of dietary fibre and starch content on digestibility, microfaunal population and caecal function was investigated in a subterranean mole-rat, Heterocephalus glaber (Rodentia). Mole-rats were fed on a diet of either sweet potato (neutral-detergent fibre (NDF) 65 g/kg dry matter (DM), starch 638 g/kg DM) or carrot (NDF 157 g/kg DM, starch 258.7 g/kg DM) for 4 weeks. Daily intake and faecal output were monitored. Thereafter caecal microfaunal population, density and function were assessed using light and scanning electron microscopy and by measuring both gas and short chain fatty acid (SCFA) production. A 2.4-fold increase in fibre and 2.5-fold decrease in starch content resulted in a decrease in caecal DM content (390 g/kg). A concomitant dramatic decline (by 93%) in ciliate protozoa with a corresponding 2-fold increase in bacteria also accompanied this change in diet. Fermentative efficiency as indicated by gas production was 2.6 times greater on a carrot diet than on sweet potato. Microbial fermentation resulted in higher SCFA concentrations on the carrot diet, with a 42 % reduction in SCFA concentration on the sweet potato diet. Here, SCFA contributed 5.1 % of daily energy expenditure and this increased 5.0-fold on the carrot diet. Caecal micro-organism function, therefore, played an important role in the nutritional physiology of these naked mole-rats, and enabled maximum utilization of the food substrate.