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The quality and quantity of light changes significantly over the course of the day. The effect of light intensity on physiological and behavioural responses of animals has been well documented, particularly during the scotophase, but the effect of the wavelength of light, particularly during the photophase, less so. We assessed the daily responses in urine production, urinary 6-sulfatoxymelatonin (6-SMT) and glucocorticoid metabolite (uGCM) concentrations in the nocturnal Namaqua rock mouse (Micaelamys namaquensis) and diurnal four striped field mouse (Rhabdomys pumilio) under varying wavelengths of near monochromatic photophase (daytime) lighting. Animals were exposed to a short-wavelength light cycle (SWLC; ∼465-470 nm), a medium-wavelength light cycle (MWLC; ∼515-520 nm) and a long-wavelength light cycle (LWLC; ∼625-630 nm). The SWLC significantly attenuated mean daily urine production rates and the mean daily levels of urinary 6-SMT and of uGCM were inversely correlated with wavelength in both species. The presence of the SWLC greatly augmented overall daily 6-SMT levels, and simultaneously led to the highest uGCM concentrations in both species. In M. namaquensis, the urine production rate and urinary 6-SMT concentrations were significantly higher during the scotophase compared to the photophase under the SWLC and MWLC, whereas the uGCM concentrations were significantly higher during the scotophase under all WLCs. In R. pumilio, the urine production rate and uGCM were significantly higher during the scotophase of the SWLC, not the MWLC and LWLC. Our results illustrate that wavelength in the photophase plays a central role in the entrainment of rhythms in diurnal and nocturnal African rodent species., (Copyright © 2021 Elsevier Inc. All rights reserved.)

The importance of haemotropic Mycoplasma (haemoplasma) infections to animal and human health is increasingly recognised. Although wild rodents are known to host these bacteria, haemoplasma prevalence and diversity in small mammals is under-documented, globally. This is due to the reliance on molecular approaches to detect these unculturable, obligate bacteria and to a paucity of assays targeting informative gene regions. We attempted to address these challenges by evaluating the performance of three 16S rRNA PCR assays for detecting Mycoplasma in four African mole-rat species of the family Bathyergidae. This was achieved by screening DNA samples prepared from lung and liver samples of 260 bathyergids, sampled from natural and urban landscapes in the Western Cape Province with one published and two novel conventional PCR assays. Sequence-confirmed Mycoplasma presence guided calculations of the relative sensitivity and specificity of the assays and revealed that 26.5% of the rodents were haemoplasma-positive. Bathyergus suillus sampled near an informal human settlement had a significantly higher infection rate (42%) than the three bathyergid species sampled from natural settings, for which PCR-positivity ranged from 0% to 36%. The 16S rRNA gene phylogeny identified the presence of six Mycoplasma strains in bathyergids that form a novel monophyletic lineage belonging to the haemofelis group, with 16S rRNA and Rnase P gene phylogenies indicating that the bathyergid-associated haemoplasmas were novel and closely related to Mycoplasma coccoides. Assay sensitivity ranged from 60.3% to 76.8% and specificity from 94.8% to 100% and both were highest for the novel assay targeting a ~ 300 bp region of the 16S rRNA gene. Results confirm the presence of novel haemoplasma strains in bathyergid species from South Africa and emphasise the need for expanded studies on haemoplama prevalence, diversity, and transmission routes in other small mammal species from this biodiverse region., (Copyright © 2021 Elsevier B.V. All rights reserved.)

Non-invasive methods for measuring glucocorticoids and their metabolites are frequently used in ecological, behavioural and physiological studies of mammals. Using faeces, urine and other matrices for such a measurement has considerable advantages in comparison to more traditional methods, but also requires thorough validation of the methods used. Eastern rock sengis (Elephantulus myurus) are fascinating African mammals and the non-invasive monitoring of the adrenocortical activity opens up new opportunities to study their biology. We were able to validate two assays for measuring urinary (uGCM) and faecal glucocorticoid metabolite (fGCM) concentrations in this species using a dose-dependent challenge with adrenocorticotropic hormone (ACTH). A higher concentration of ACTH elicited higher uGCM and fGCM concentrations in both males and females. Interestingly, uGCM and fGCM concentrations and the responses to ACTH were higher in females than in males and small changes in faecal glucocorticoid metabolites could not be reliably detected in males. In contrast to ACTH, a saline injection did not result in an increase in uGCM or fGCM concentrations. The study also provided insight into when responses to a stressor are likely to be detected in the urine and faeces of sengis and opens up new opportunities to study the stress physiology of this and other sengi species. It further emphasises the importance of thoroughly validating non-invasive methods for measuring hormones in both sexes of a species and for incorporating dose-dependent approaches., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2020 Elsevier Inc. All rights reserved.)

Bartonellosis is a vector-borne disease that is often misdiagnosed due to a broad range of clinical symptoms, compounded by a lack of awareness regarding the prevalence, diversity and public health impacts of regional strains. Despite recent PCR-based confirmation of Bartonella in 9.7% of non-malarial, acute febrile patients in South Africa, data regarding reservoirs of infection are limited. As the majority of Bartonella species described to date are associated with rodent species globally, including zoonotic species such as B. elizabethae, and as rodent biodiversity is high in southern Africa, we evaluated Bartonella in the Tete Veld rat (Aethomys ineptus), a highly adaptable murid rodent that thrives in both natural and commensal settings. These rodents are infested with a broad range of ectoparasite species, and often occur in sympatry with Micaelamys namaquensis, an indigenous rodent previously shown to host B. elizabethae. DNA extracts from heart samples of 75 A. ineptus trapped over an eight-month period, from the Roodeplaat Nature Reserve (RNR), were evaluated using a multi-locus sequence analysis (MLSA) approach. Nucleotide sequencing and phylogenetic analyses of individual (gltA, ribC, rpoB and nuoG) and concatenated gene datasets confirmed the presence of three discrete Bartonella lineages (I-III). Lineages I and II, are genetically distinct from all currently recognised Bartonella species but cluster with strains present in other indigenous rodents from South and East Africa, whereas lineage III contained B. elizabethae, a zoonotic species associated with Rattus species globally. Records confirming R. tanezumi presence in this nature reserve, which is situated in close proximity to Pretoria, the administrative capital of South Africa, suggests the likelihood of spill-over from invasive to indigenous species. These results together with the high levels of infection (86.7%) and co-infection (33.8%), indicate that A. ineptus is a natural reservoir for multiple Bartonella species in South Africa, including one with zoonotic potential., (Copyright © 2019 Elsevier B.V. All rights reserved.)

Bartonellae bacteria are associated with several re-emerging human diseases. These vector-borne pathogens have a global distribution, yet data on Bartonella prevalence and diversity in the Arabian Peninsula are limited. In this study we assessed the Bartonella infection status of the Baluchistan gerbil (Gerbillus nanus), a species associated with pastoral communities throughout the Middle East region, using a multi-gene PCR screening approach. The results demonstrated that 94 (68.1%) of the 138 gerbils trapped on a monthly basis, over a period of one year, were PCR-positive. Sequencing of the gltA gene region confirmed the presence of four discrete Bartonella lineages (I-IV) and high levels of co-infection (33.0%). Each of the four lineages, varied in overall abundance (7.5%-47.9%) and had discernible seasonal peaks. Bartonella status was significantly correlated with ectoparasite presence, but not with sex, nor with season. Statistical analyses further revealed that co-infected individuals had a significantly higher relative body condition. Multi-locus sequence analysis (MLSA) performed with a concatenated dataset of three genetic loci (gltA, nuoG, and rpoB), 1452 nucleotides (nt) in length confirmed that lineage IV, which occurred in 24 PCR-positive animals (25.5%), is most closely related to zoonotic B. elizabethae. The remaining three lineages (I-III) formed a monophyletic clade which, on the basis of gltA was shown to contain bartonellae from diverse Gerbillinae species from the Middle East, suggestive of a gerbil-associated species complex in this region. Lineage I was identical to a Candidatus B. sanaae strain identified previously in Bushy-tailed jirds (Sekeetamys calurus) from Egypt, wherease MLSA indicate that lineages II and III are novel. The high levels of infection and co-infection, together with the presence of multiple Bartonella lineages indicate that Gerbillus nanus is likely a natural reservoir of Bartonella in the Arabian Peninsula., (Copyright © 2018 Elsevier B.V. All rights reserved.)

In Damaraland mole-rats (Fukomys damarensis), non-breeding subordinates contribute to the care of offspring born to the breeding pair in their group by carrying and retrieving young to the nest. In social mole-rats and some cooperative breeders, dominant females show unusually high testosterone levels and it has been suggested that high testosterone levels may increase reproductive and aggressive behavior and reduce investment in allo-parental and parental care, generating age and state-dependent variation in behavior. Here we show that, in Damaraland mole-rats, allo-parental care in males and females is unaffected by experimental increases in testosterone levels. Pup carrying decreases with age of the non-breeding helper while the change in social status from non-breeder to breeder has contrasting effects in the two sexes. Female breeders were more likely than female non-breeders to carry pups but male breeders were less likely to carry pups than male non-breeders, increasing the sex bias in parental care compared to allo-parental care. Our results indicate that testosterone is unlikely to be an important regulator of allo-parental care in mole-rats., (Copyright © 2018. Published by Elsevier Inc.)

The Arabian oryx, a moderately large mammal that inhabits a harsh desert environment, has been shown to exhibit seasonal variations in activity and inactivity patterns. Here we analyzed the continuous year-round activity patterns of twelve free-roaming Arabian oryx under natural conditions from two varying desert environments in Saudi Arabia using abdominally implanted activity meters. We simultaneously recorded weather parameters at both sites to determine whether environmental factors are responsible for temporal niche switching as well as the seasonal structuring and timing of this behavioural plasticity. Our results demonstrate that Arabian oryx undergo temporal niche switching of 24h activity patterns at a seasonal level and exhibit distinct nocturnal/crepuscular activity during summer, diurnal activity during winter and intermittent patterns of behaviour during the transitional seasons of autumn and spring. In addition, the oryx exhibited inter- and intra-seasonal variations in the temporal budgeting of 24h activity patterns. Strong relationships with both photoperiod and ambient temperatures were found and in some instances suggested that increasing ambient temperatures are a primary driving force behind seasonal shifts in activity patterns. These adaptive patterns may be dictated by the availability of food and water, which in turn are strongly influenced by seasonal climate variations. Overall, the adaptive responses of free-roaming Arabian oryx in such harsh and non-laboratorial conditions provide a framework for comparing wild populations as well as aiding conservation efforts., (Copyright © 2017 Elsevier Inc. All rights reserved.)

A previous study investigating potential adult hippocampal neurogenesis in microchiropteran bats failed to reveal a strong presence of this neural trait. As microchiropterans have a high field metabolic rate and a small body mass, it is possible that capture/handling stress may lead to a decrease in the detectable presence of adult hippocampal neurogenesis. Here we looked for evidence of adult hippocampal neurogenesis using immunohistochemical techniques for the endogenous marker doublecortin (DCX) in 10 species of microchiropterans euthanized and perfusion fixed at specific time points following capture. Our results reveal that when euthanized and perfused within 15 min of capture, abundant putative adult hippocampal neurogenesis could be detected using DCX immunohistochemistry. Between 15 and 30 min post-capture, the detectable levels of DCX dropped dramatically and after 30 min post-capture, immunohistochemistry for DCX could not reveal any significant evidence of putative adult hippocampal neurogenesis. Thus, as with all other mammals studied to date apart from cetaceans, bats, including both microchiropterans and megachiropterans, appear to exhibit substantial levels of adult hippocampal neurogenesis. The present study underscores the concept that, as with laboratory experiments, studies conducted on wild-caught animals need to be cognizant of the fact that acute stress (capture/handling) may induce major changes in the appearance of specific neural traits., (Copyright © 2014 IBRO. Published by Elsevier Ltd. All rights reserved.)

In the present study, orexinergic cell bodies within the brains of rhythmic and arrhythmic circadian chronotypes from three species of African mole rat (Highveld mole rat-Cryptomys hottentotus pretoriae, Ansell's mole rat--Fukomys anselli and the Damaraland mole rat--Fukomys damarensis) were identified using immunohistochemistry for orexin-A. Immunopositive orexinergic (Orx+) cell bodies were stereologically assessed and absolute numbers of orexinergic cell bodies were determined for the distinct circadian chronotypes of each species of mole rat examined. The aim of the study was to investigate whether the absolute numbers of identified orexinergic neurons differs between distinct circadian chronotypes with the hypothesis of elevated hypothalamic orexinergic neurons in the arrhythmic chronotypes compared with the rhythmic chronotypes. We found statistically significant differences between the circadian chronotypes ofF. anselli, where the arrhythmic group had higher mean numbers of hypothalamic orexin neurons compared with the rhythmic group. These differences were observed when the raw data was compared and when the raw data was corrected for body mass (M(b)) and brain mass (M(br)). For the two other species investigated, no significant differences were noted between the chronotypes, although a statistically significant difference was noted between all rhythmic and arrhythmic individuals of the current study when the counts of orexin neurons were corrected for M(b)--the arrhythmic individuals had larger numbers of orexin cells., (Copyright © 2011 IBRO. Published by Elsevier Ltd. All rights reserved.)

Spatial learning and memory is an important skill for the survival and fitness and may vary between the sexes depending on differences in space use. This is particularly true for animals that explore the subterranean niche as it is associated with high travelling costs. In subterranean rodents the complexity of burrow systems varies with differing degrees of sociality possibly posing stronger selective pressures regarding spatial abilities on species with more complex burrow structures. This could lead to superior abilities in spatial learning and memory in social compared to solitary subterranean species. We tested this hypothesis in two species of subterranean mole-rats, the eusocial Damaraland (Fukomys damarensis) and solitary Cape mole-rats (Georychus capensis) by comparing their ability to locate food in an artificial maze. Measurements of the time taken to the goal chamber, the number of wrong turns taken, and the average velocity at which animals travelled were used to compare performance between animals. We did not find marked sex-specific differences in either study species during the assessment of learning and memory retention. In accordance with our hypothesis significant differences between the species were apparent during both learning and memory trials with the social species exhibiting superior performances. However, in both species memory retention was generally high suggesting that the fossorial lifestyle poses a strong selective pressure on spatial abilities in subterranean mammals.

Mole-rat species within the family Bathyergidae exhibit a wide range of reproductive strategies and social systems. Various forms of reproductive suppression are displayed within this family: in the solitary species, breeding is suspended for part of the year and in the social species, reproduction is suppressed in subordinate animals. This study investigated the gonadotrophin-releasing hormone 1 (GnHR-1) systems of breeding and non-breeding solitary Cape mole-rats and social Natal mole-rats for possible inter- and/or intra-species differences. In both species, GnRH-1 cell bodies are predominantly in the medial septum region of the diagonal band or the preoptic area, with relatively few in the mediobasal hypothalamus; a dense concentration of GnRH-1-immunoreactive (ir) processes is present in the region of the organum vasculosum of the lamina terminalis. In Cape mole-rats, GnRH-1-ir processes are particularly dense within the lateral margins of the median eminence, which is enfolded by a large pars tuberalis of the pituitary gland. Natal mole-rats display GnRH-1-ir processes across the breadth of the median eminence, which is abutted by a relatively small pars tuberalis. There are more GnRH-1-ir cell bodies in Natal mole-rats than in Cape mole-rats ( approximately 720 vs. approximately 420). No significant differences were found in the number, distribution or size of GnRH-1-ir cell bodies according to season in Cape mole-rats or according to reproductive status or sex in Natal mole-rats. In female and male Natal mole-rats, GnRH-1-immunoreactivity in the median eminence is less dense in the reproductive animals; no such difference was found in Cape mole-rats between the breeding and non-breeding seasons. These immunohistochemical results are discussed in the light of earlier studies which identified no functional neuroendocrine impediments underlying regulated reproduction in either Cape or Natal mole-rats. The cumulative findings suggest that the principal factors determining seasonal or socially induced suppression of reproduction in these species are behavioral rather than neuroendocrine.

The gonadotrophin-releasing hormone 1 (GnRH1) system has been investigated immunohistochemically in Cape dune mole-rats (Bathyergus suillus), subterranean rodents that normally display severe aggression towards conspecifics. These animals breed seasonally and show a reduced mean plasma level of luteinising hormone during the non-breeding season. GnRH1-immunoreactive (ir) cell bodies and processes are found in the septal/preoptic area and the mediobasal hypothalamus; the cell bodies are found in equal measure in these two regions. Dense aggregations of GnRH1-ir fibres are present in the organum vasculosum of the lamina terminalis and the external zone of the median eminence. The total number of detectable GnRH1-ir cell bodies does not differ between the sexes or within the sexes between breeding and non-breeding seasons. Similarly there is no difference in the distribution of detectable GnRH1-ir cell bodies in male and female mole-rats in and out of the breeding season. Although the average size of GnRH1-ir cell bodies does not differ between the seasons in males, their size in females is significantly smaller in the non-breeding season. Whether this reduced size reflects reduced GnRH1 synthesis remains to be determined.

In some animal societies, males vary in the strategies and tactics that they use for reproduction. Explanations for the evolution of alternative tactics have usually focussed on extrinsic factors such as social status, the environment or population density and have rarely examined proximate differences between individuals. Anecdotal evidence suggests that two alternative reproductive tactics occur in cooperatively breeding male Cape ground squirrels. Here we show that there is strong empirical support for physiological and behavioural differences to uphold this claim. 'Dispersed' males have higher resting metabolic rates and a heightened pituitary activity, compared with philopatric 'natal' males that have higher circulating cortisol levels. Dispersed males also spend more time moving and less time feeding than natal males. Additionally, lone males spend a greater proportion of their time vigilant and less of their time foraging than those that were in groups. The choice of whether to stay natal or become a disperser may depend on a number of factors such as age, natal group kin structure and reproductive suppression, and the likelihood of successful reproduction whilst remaining natal. Measuring proximate factors, such as behavioural and endocrine function, may provide valuable insights into mechanisms that underlie the evolution of alternative reproductive tactics.

We have studied the visual system of subterranean mole-rats of the rodent family Bathyergidae, for which light and vision seem of little importance. The eye diameter varies between 3.5mm in Bathyergus suillus and 1.3mm in Heterocephalus glaber. The small superficial eyes have features typical of sighted animals (clear optics, well-developed pupil and well-organized retina) and appear suited for proper image formation. The retinae are rod-dominated but possess rather high cone proportions of about 10%. The total number of retinal ganglion cells and optic nerve fibres ranges between 6000 in Bathyergus suillus and 2100 in Heliophobius argenteocinereus. Visual acuity (estimated from counts of peak ganglion cell density and axial length of the eye) is low, ranging between 0.3 and 0.5 cycles/degree. The retina projects to all the visual structures described in surface-dwelling sighted rodents. The suprachiasmatic nucleus is large and receives bilateral retinal input. All other visual nuclei are reduced in size and receive almost exclusively contralateral retinal projections of varying magnitude. The primary visual cortex is small and, in comparison to other rodents, displaced laterally. In conclusion, the African mole-rats possess relatively well-developed functional visual subsystems involved in photoperiodicity, form and brightness discrimination. In contrast, visual subsystems involved in coordination of visuomotor reflexes are severely reduced. This pattern suggests the retention of basic visual capabilities. Residual vision may enable subterranean mammals to localize breaches in the burrows that let in light thus providing a cue to enable mole-rats to reseal such entry points and to prevent entry of predators.

Small rodents with a large surface-area-to-volume ratio and a high thermal conductance are likely to experience conditions where they have to expend large amounts of energy in order to maintain a constant body temperature at low ambient temperatures. The survival of small rodents is thus dependent on their ability to reduce heat loss and increase heat production at low ambient temperatures. Two such animals are the social subterranean rodents Cryptomys damarensis (the Damaraland mole-rat) and Cryptomys hottentotus natalensis (the Natal mole-rat). This study examined the energy savings associated with huddling as a behavioural thermoregulatory mechanism to conserve energy in both these species. Individual oxygen consumption (VO(2)) was measured in groups ranging in size from one to 15 huddling animals for both species at ambient temperatures of 14, 18, 22, 26 and 30 degrees C. Savings in energy (VO(2)) were then compared between the two species. Significant differences in VO(2) (p<0.05) were found within each species, indicating that both Damaraland mole-rats and Natal mole-rats saved more energy in larger as opposed to smaller groups. VO(2) was also different between the two species, with Damaraland mole-rats showing a higher decrease in VO(2) with increasing group size compared to Natal mole-rats. These findings suggest that huddling confers significant energy savings in both species and that the amount of energy saved is related to each species' ecology. More generally, these findings suggest that group living desert-adapted species are likely to be more prone to heat loss at low ambient temperatures than temperate-adapted species, especially at low group sizes. This is presumably offset against the advantages obtained by having a low metabolic rate and avoiding hyperthermia when temperatures are hot.

Subterranean mammals (those that live and forage underground) inhabit a challenging microenvironment, with high levels of carbon dioxide and low levels of oxygen. Consequently, they have evolved specialised morphological and physiological adaptations. For small mammals that inhabit high altitudes, the effects of cold are compounded by low oxygen partial pressures. Hence, subterranean mammals living at high altitudes are faced with a uniquely demanding physiological environment, which presumably necessitates additional physiological adjustments. We examined the thermoregulatory capabilities of two populations of Lesotho mole-rat Cryptomys hottentotus mahali that inhabit a 'low' (1600 m) and a 'high' (3200 m) altitude. Mole-rats from the high altitude had a lower temperature of the lower critical point, a broader thermoneutral zone, a lower thermal conductance and greater regulatory non-shivering thermogenesis than animals from the lower altitude. However, minimum resting metabolic rate values were not significantly different between the populations and were low compared with allometric predictions. We suggest that thermoregulatory costs may in part be met by animals maintaining a low resting metabolic rate. High-altitude animals may adjust to their cooler, more oxygen-deficient environment by having an increased non-shivering thermogenesis whilst maintaining low thermal conductance.

Previous studies have suggested that nocturnal and diurnal species of rodents differ in their circadian responses to light including phase shifts and early gene expression. Rhabdomys pumilio, the four-striped field mouse, is diurnal both in nature and in the laboratory. We studied in this species the response of the suprachiasmatic nucleus (SCN) to light stimuli at different time periods using light-induced expression of Fos as marker of neuronal activity. Fos induction in the SCN was investigated using immunohistochemistry and quantitative image analysis. The animals were exposed to a 15 min light pulse with monochromatic green light at different circadian times throughout a 24-h cycle. Animals maintained in constant darkness served as controls. R. pumilio exhibited an endogenous Fos rhythm in the SCN during constant darkness with highest expression during the subjective day at circadian time (CT) 2 and CT10. Photic stimulation resulted in significant Fos induction in the SCN at CT6, CT14, CT18 and CT22, compared to controls kept in constant darkness, with a peak of expression at CT22, i.e. during late subjective night, mainly due to expression in the ventral SCN. In tract tracing experiments based on the use of cholera toxin subunit B, we found that retinal fibres innervate mainly the contralateral ventral SCN. The intergeniculate leaflet received bilateral retinal innervation with overlap between ipsilateral and contralateral fibres. Altogether the data show that the rodent R. pumilio is a unique diurnal model for chronobiological studies.

The Lesotho mole-rat is a social subterranean rodent that occurs at altitude in the Drakensberg mountain range. As a consequence of living permanently underground these animals rarely if ever are exposed to light. The visual system of African mole-rats is particularly regressed whereas the circadian system is proportionately conserved. This study investigated the locomotor activity patterns of 12 Lesotho mole-rats maintained under a range of different lighting regimes. The majority (91.7%) of mole-rats entrained their activity patterns to a LD photoperiod of 12L/12D. The mole-rats displayed a monophasic nocturnal activity preference. Under constant dark (DD) most of the mole-rats (83.3%) showed a free running circadian activity pattern with a tau of 23.8 h to 24.4 h (mean+/-S.E.M.: 24.07 h+/-0.07 h; n=10). The phase of the activity rhythms each mole-rat exerted during the previous LD-cycle did not change when the animals started free-running after being placed in constant conditions. The duration of re-entrainment to a second bout of LD 12:12 amounted to 9.4+/-2.03 days (mean+/-S.E.M., n=10). Eleven mole-rats (91.7%) adjusted their locomotor activity rhythms to an inversed light regime DL 12:12 and displayed significant nocturnal activity preference. The animals required 9.73+/-2.01 days (mean+/-S.E.M., n=11) to adjust to the DL-photoperiod. The Lesotho mole-rat thus possesses a functional circadian clock that responds to a photic zeitgeber.

In this paper we investigated the effect of sex and season on baseline and post-GnRH challenge luteinising hormone (LH) levels in a solitary, seasonally breeding mole-rat. Circulating basal concentrations of luteinising hormone (LH) were found to differ significantly with season in both sexes. However, no significant difference was found in circulating basal LH concentration between the sexes either within or out of the breeding season. The magnitude of the LH response to an exogenous pharmacological overdose of GnRH both in and out of the breeding season in males and females respectively was not significant. This finding suggests that there is no down regulation of GnRH receptors on the pituitary during the non-breeding season. Cape dune mole-rats thus have the potential for opportunistic breeding outside of the typical breeding period. We argue that this represents an adaptation to limited and brief opportunities for mating in this xenophobic and aggressive species.

Do burrowing mammals that naturally experience hypoxic and hypercapnic conditions exhibit modifications to the blood chemistry at high altitudes? We investigated two populations of the Lesotho mole-rat living at different altitudes in the highlands of the Drakensberg. There was no significant difference between the specimens from 3200 and 1600 m in mean red blood cell count (RCC=8.9x10(6)+/-1.6x10(6) vs. 8.4x10(6)+/-0.95x10(6) mm3, respectively) or packed red cell volumes (haematocrit=0.51+/-0.06 vs. 0.49+/-0.05, respectively). However, blood haemoglobin (Hb) concentration was significantly higher in the high altitude than in the low-altitude specimens (178+/-9 vs. 160+/-16 g/l). The oxygen equilibrium curves of thawed whole blood showed no displacement to the left in the animals sampled at the higher elevation. The data indicate that the oxygen-transporting properties of mole-rat blood do not change markedly with increased elevation and that burrowing mammals are a priori hypoxia-adapted.

The influence of endogenous opioid peptides (EOPs) on plasma luteinizing hormone (LH) levels in subordinate female highveld mole-rats (Cryptomys hottentotus pretoriae) was investigated to elucidate the physiological mechanisms responsible for inhibiting their gonadotropin-releasing hormone (GnRH) and/or LH release. The opioid antagonist naloxone was administered either alone or with GnRH. A single injection of naloxone failed to alter plasma LH levels in dominant reproductive females or in subordinate non-reproductive females in the presence or absence of their ovaries. Pituitary sensitivity to a GnRH challenge was not influenced by naloxone administered acutely or according to longer-term regimens in any of the treatment groups. The results suggest no role for EOPs at the level of the pituitary or hypothalamus in the socially induced infertility evident in non-reproductive female highveld mole-rats.

The Damaraland mole-rat is a eusocial, subterranean rodent that exhibits a seasonal breeding. Non-reproductive females show physiological suppression of reproduction whilst in the confines of the natal colony. This study set out to investigate whether dispersing female Damaraland mole-rats exhibit induced or spontaneous ovulation. Fifteen non-reproductive females were removed from their natal colonies and housed individually for a period of 6 weeks. During this period urine was collected from all animals every second day. After this initial period the animals were divided into 3 groups. Females were subjected to 1 of 3 trials: a control group housed separately without a male, allowed non-physical contact, or placed in direct physical contact with vasectomized males. Urine was collected for a further 5 weeks, and urinary progesterone profiles established. All three groups showed a significant difference in urinary progesterone concentrations between the two treatment periods indicating initiation of follicular development in all animals following removal from the colony. Histological results further revealed that at least one animal in the control group and five of the six animals in the vasectomized group had corpora lutea present in the ovaries showing that ovulation has taken place. All groups had similar numbers of Graafian follicles, indicating that all females were likely to ovulate in the near future. This finding suggests that females are capable of spontaneous ovulation, but the act of coitus may advance the onset of ovulation in this arid dwelling mole-rat.

Although humans are diurnal in behaviour, animal models used for the study of circadian rhythms are mainly restricted to nocturnal rodents. This study focussed on the circadian behaviour of a rodent from South Africa that has a preference for daylight, the four-striped field mouse, Rhabdomys pumilio. In order to characterise the behavioural pattern of daily activity, locomotor rhythms were studied under different light regimes using an automated data recording system. Under conditions of natural daylight, which include dawn and dusk transitions, R. pumilio exhibited activity restricted to the daytime period. Activity was concentrated around morning and evening with a decrease during mid-day. A similar diurnal preference pattern of behaviour was recorded under a light-dark cycle of artificial illumination. Under conditions of constant darkness, the four-striped field mouse exhibited a free-running circadian rhythm of locomotor activity with activity concentrated during the subjective day. Free-running rhythms varied greatly between individuals, from slightly less to slightly more than 24 h (range = 23.10 to 24.80 h). Under conditions of constant light, the mice were more active during subjective day, but the free-running rhythm in all individuals was consistently longer than 24 h (range = 24.30 to 24.79 h).

Winter is an energetically stressful period for small mammals as increasing demands for thermoregulation are often coupled with shortages of food supply. In sub-tropical savannah, Hottentot golden moles (Ambysomus hottentottus longiceps) forage throughout the year and for long periods of each day. This may enable them to acquire sufficient resources from an insectivorous prey base that is both widely dispersed and energetically costly to obtain. However, they also inhabit much cooler regions; how their energy budgets are managed in these areas is unknown. We measured the daily energy expenditure (DEE), resting metabolic rate (RMR) and water turnover (WTO) of free-living golden moles during both winter and summer at high altitude (1500 m). We used measurements of deuterium dilution to estimate body fat during these two periods. DEE, WTO and body mass did not differ significantly between seasons. However, RMR values were higher during the winter than the summer and, in the latter case were also lower than allometric predictions. Body fat was also higher during the winter. Calculations show that during the winter they may restrict activity to shorter, more intense periods. This, together with an increase in thermal insulation, might enable them to survive the cold.

The Mashona mole rat, Cryptomys darlingi, is a social, subterranean African rodent that is rarely, if ever, exposed to light, and that exhibits a regressed visual system. This study investigated locomotor activity patterns of Mashona mole rats (n=12) under different light cycles. Activity was measured using either infrared captors (n=8) or running wheels (n=4). The mole rats entrained their activity to a standard (LD 12:12) photoperiod. They displayed either a nocturnal or diurnal activity preference with one bout of activity and one bout of rest. Therefore, as a species, the Mashona mole rat did not show a clear nocturnal or diurnal activity preference. When the LD (12:12) light cycle was inversed, the animals switched their activity, too. Under constant dark (DD), most mole rats (73%) showed a free-running circadian activity rhythm, but under constant light (LL), only some (36%) did. The free-run period of the rhythm (tau) ranged from 23.83 to 24.10 h. The remaining animals were arrhythmic. There was large interindividual and intraindividual variations in the rate and extent of entrainment, time of activity preference, and activity patterns. Possible reasons for the observed variations are discussed. It is concluded that the Mashona mole rat has an endogenous activity rhythm which approximates 24 h, that the mole rat can distinguish between light and dark, and that the endogenous clock utilises this photic information as a zeitgeber.

The hormone melatonin is synthesised and secreted from the pineal gland in darkness and triggers the daily and seasonal timing of various physiological and behavioural processes. The Mashona mole-rat, Cryptomys darlingi, lives in subterranean burrows that are completely sealed and is therefore rarely, if ever, exposed to light under natural conditions. Hence, this species is of particular interest for studies on rhythms of melatonin secretion. We investigated how plasma melatonin concentrations of the Mashona mole-rat responded to exposure to a long-term standard photoperiod of 12 h light, 12 h dark (12:12 LD), constant light (LL) and constant dark (DD). In addition, we examined whether plasma melatonin concentration was coupled to locomotor activity. Mashona mole-rats displayed rhythms of plasma melatonin concentration that appeared entrained to the standard LD photoperiod, suggesting that the mole-rat is capable of perceiving and entraining to this photic zeitgeber. Furthermore, under chronic constant lighting conditions (DD, LL), circadian rhythms in plasma melatonin concentration were observed, suggesting the possible existence of an endogenous rhythm. Light suppressed melatonin secretion, but constant light did not abolish the rhythm of plasma melatonin concentration. Between active and non-active animals, no difference in plasma melatonin concentration was found for any of the sequential photoperiods (LD1 DD, LD2, LL), tentatively suggesting that the rhythm of melatonin secretion is uncoupled from that of locomotor activity.

Mole-rats are strictly subterranean rodents that are rarely exposed to environmental light. They are well adapted to their environment and have reduced eyes and a severely regressed visual system. It has been shown, however, that mole-rats do exhibit endogenous circadian rhythms that can be entrained, suggesting an intact and functional circadian system. To determine whether light is the entraining agent in these animals, Fos expression in response to light pulses at different circadian times was investigated to obtain phase response curves. Light is integrated effectively in the suprachiasmatic nucleus of the Cape mole-rat (Georychus capensis), and Fos expression is gated according to the phase of the circadian clock. The Fos response in the Cape mole-rat was comparable to that of aboveground rodents. In contrast, the highveld mole-rat (Cryptomys hottentotus pretoriae) was less sensitive to light and did not show a selective Fos response according to the phase of the circadian cycle. Social species appear to be less sensitive to light than their solitary counterparts, which compares well with results from locomotor activity studies.

The Natal mole-rat, Cryptomys hottentotus natalensis, rarely, if ever, is exposed to external light cues because it occurs in completely sealed tunnel systems. As a result, their classical visual system is regressed, and therefore, their circadian system is expected proportionally to be expanded. Locomotor activity was investigated under a number of different photic regimes. Nine of the 12 mole-rats exhibited endogenous circadian rhythms of locomotor activity under constant darkness, with a mean free run period of 24.13 h (range 23.93-24.13 h), with these animals entrained to a light-dark cycle (12 L:12 D). Because C. hottentotus natalensis are able to entrain their locomotor activity to an external light source, light must reach the suprachiasmatic nucleus (SCN), suggesting a functional circadian clock. A clear day-night rhythm of melatonin secretion in animals housed under a neutral photoperiod (12 L:12 D) was observed, with higher melatonin concentrations in the dark compared with the light phase. The rhythm was maintained after the animals were transferred to either continuous light (LL) or dark (DD), suggesting that the endogenous rhythm was maintained under acute exposure to light and dark. However, under DD, the rhythm appeared to shift slightly, potentially as a result of the rhythm free running. These results show that C. hottentotus natalensis has endogenous rhythms of both locomotor activity and melatonin secretion, which are modulated by light.

Intolerance to familiar conspecifics characterises solitary mole-rats and distinguishes them from social ones. However, no study has compared the patterns of tolerance to unfamiliar conspecifics. Theoretically, both solitary and social species should react similarly and show intolerance to unfamiliar same-sex conspecifics, unless the evolution of grouping has favoured higher tolerance to conspecifics among the social species. Our study compares tolerance to unfamiliar conspecifics in four African mole-rat species exhibiting varying degrees of sociality. Dyadic encounters between female unfamiliar conspecifics were performed in a neutral arena, and the assessment of social tolerance was based on both behavioural observations of amicable contact behaviour as opposed to aggression and avoidance behaviour and the assessment of a stress response to the encounter measured as an increase in plasma cortisol concentrations. Our results show that the two highly social species and the solitary one presented similar high levels of agonistic behaviours during encounters with unfamiliar conspecifics. Nevertheless, all three social species displayed social tolerance and did not show a stress arousal during encounters with unfamiliar conspecifics, a pattern that contrasted significantly with that evidenced in the solitary species. The results suggest that physiological and behavioural characteristics allow a higher tolerance to unfamiliar conspecifics in the social as opposed to the solitary mole-rat, and the adaptive value of these characteristics are discussed. Finally, we discuss why constraints on social tolerance may be an important limiting factor to take into account in theories concerning the evolution of grouping.