Search

Genetic adaptation is the change of a population toward a phenotype that best fits the present ecological conditions of the environment it inhabits. As environmental conditions change, allele frequencies shift, resulting in different populations of the same species possessing genetic variation and divergent phenotypes. Cooperatively breeding common mole-rats (Cryptomys hottentotus hottentotus) inhabit environments along an aridity gradient in South Africa, which provides an opportunity for local genetic adaptations to occur. Using one mitochondrial gene (cytochrome b) and 3,540 SNP loci across the whole genome, we determined the phylogenetic relationship, population structure and genetic diversity of five populations of C. h. hottentotus located along an aridity gradient. Mitochondrial data identified population-specific clades that were less distinct in the two mesic populations, potentially indicating historical or recent gene flow, or the retention of ancestral haplotypes. Arid and semi-arid populations formed a distinct cluster from the non-arid populations. Genetic diversity and gene flow were higher in arid-dwelling individuals, suggesting greater connectivity and interactions between colonies in arid regions in comparison to mesic ones. Using an Aridity Index, we determined that isolation by environment, rather than isolation by geographical distance, best explains the genetic distance between the populations. Further analyses using target loci may determine if there are differing underlying genetic adaptations among populations of C. h. hottentotus. These analyses could help unravel population differences in response to environmental factors within a subspecies of bathyergid mole-rat and determine the adaptive capacity of this small nonmigratory subterranean rodent species in response to aridification in the face of climate change. [ABSTRACT FROM AUTHOR]

Sperm competition results from postcopulatory continuation of male–male competition for paternity. The level of sperm competition is predicted to be highest in species with greater polyandry and weakest in monogamous pairs. Sperm competition levels can be indexed using traits that reflect male investment in fertilization, particularly relative testes mass (RTM). However, the relationship between RTM and levels of sperm competition may also be influenced by precopulatory competition selecting for higher levels of testosterone, also produced by the testes. To test the relationship between RTM and both pre- and postcopulatory male–male competition we compared two bathyergid mole-rat species, the promiscuous Georychus capensis and the monogamous eusocial Fukomys damarensis. The promiscuous species had not only larger RTM, but also a greater proportion of spermatogenic tissue, maximizing germ cell production as well. Conversely, the eusocial species had smaller testes, but a higher proportion of interstitial tissue (which contains the androgenic Leydig cells) and higher levels of testosterone. Consequently, testicular traits as well as testes mass may be under selection, but these are not normally measured. More research is required on relative investment in different testicular traits in relation to both pre- and postcopulatory selection pressures. [ABSTRACT FROM AUTHOR]

The Mashona mole-rat, Fukomys darlingi (Thomas, 1895), is a little studied social African mole-rat (Bathyergidae) from south-astern Africa. Here, we present an integrative study characterizing the genetic diversity of populations assigned to F. darlingi with special focus on animals from Nsanje, southern Malawi. These mole-rats show pronounced differences in body mass and general appearance compared to nominate F. darlingi from Zimbabwe and Mozambique, but their taxonomic status has so far remained unclear. A genetic analysis encompassing all major lineages of the genus Fukomys suggests that this population indeed represents a deeply nested lineage within the F. darlingi clade. The karyotype of the Nsanje mole-rats also corresponds to that of the nominate form, being 2n = 54. While both nuclear and mitochondrial data agree about the assignment of the Nsanje mole-rats to F. darlingi, our analyses revealed substantial mitonuclear discordance for other branches within the Fukomys phylogenetic tree. Nsanje mole-rats are significantly larger than nominate F. darlingi and their ontogeny and reproduction closely resemble similar-sized congeneric species rather than the nominate population. The somatic growth of the Nsanje form is the slowest of all African mole-rats. The maximum life span of F. darlingi is at least 19 years. The observed differences between nominate F. darlingi and mole-rats from Nsanje may be attributed mainly to their different body mass. Our study highlights the advantages of an integrative approach for understanding the diversity of African mole-rats and emphasizes the great intraspecific variability that may be encountered in these underground-dwelling rodents. [ABSTRACT FROM AUTHOR]

Simple Summary: Naked mole-rats (Heterocephalus glaber) have unique physiological and behavioral characteristics that make them a species of interest in biomedical research. However, due to their small size, frequent blood sampling to monitor the animals is not possible. Thus, we aimed to validate enzyme immunoassays, tools which detect the quantity of hormone molecules, for monitoring stress-related hormones, glucocorticoids, and/or their metabolites, in naked mole-rats using urine and feces. We administered a high and low dose of adrenocorticotrophic hormone, which causes the animal to physiologically respond to stress, and a saline administration as a control, which should not elicit a physiological stress response. The results revealed two enzyme immunoassays, namely, a 5α-pregnane-3β,11β,21-triol-20-one detecting glucocorticoid metabolites with a 5α-3β-11β-diol structure and an 11-oxoaetiocholanolone detecting glucocorticoid metabolites with a 5β-3α-ol-11-one structure, for measuring stress-related hormones in male and female urine, respectively. Furthermore, one enzyme immunoassay, namely, 11-oxoaetiocholanolone detecting 11,17 dioxoandrostanes for measuring stress-related hormones in feces of both sexes. There were sex-related differences in how the animals responded to the high and low doses of the adrenocorticotrophic hormone. We recommend using feces to monitor stress-related hormones in naked mole-rats, which will be valuable when investigating housing conditions and other animal welfare aspects. Small size in mammals usually restricts long-term, frequent monitoring of endocrine function using plasma as a matrix. Thus, the non-invasive monitoring of hormone metabolite concentrations in excreta may provide an invaluable approach. The aim of the current study was to examine the suitability of enzyme immunoassays (EIAs) for monitoring responses to stressors in the naked mole-rat (Heterocephalus glaber, NMR) using urine and feces as hormone matrices. A saline control administration, and a high- and low-dose adrenocorticotropic hormone (ACTH) challenge were performed on six male and six female disperser morph NMRs. The results revealed that a 5α-pregnane-3β,11β,21-triol-20-one EIA detecting glucocorticoid metabolites (GCMs) with a 5α-3β-11β-diol structure is the most suitable assay for measuring concentrations in male urine samples, whereas an 11-oxoaetiocholanolone EIA detecting GCMs with a 5β-3α-ol-11-one structure appears the most suitable EIA for quantifying GCMs in female urine. An 11-oxoaetiocholanolone EIA detecting 11,17 dioxoandrostanes was the most suitable EIA for quantifying GCMs in the feces of both sexes. There were sex-related differences in response to the high- and low-dose ACTH challenge. We recommend using feces as a more suitable matrix for non-invasive GCM monitoring for NMRs which can be valuable when investigating housing conditions and other welfare aspects. [ABSTRACT FROM AUTHOR]

Published studies and ten new unpublished records included herein reveal that approximately 174 species of endoparasites (helminths and protozoans) are known from 65 of 163 species of rodents that occupy the subterranean ecotope globally. Of those, 94 endoparasite species were originally described from these rodents. A total of 282 host-parasite associations are summarized from four major zoogeographic regions including Ethiopian, Palearctic/Oriental, Nearctic, and Neotropical. Thirty-four parasite records from the literature have been identified to only the level of the genus. In this summary, ten new records have been added, and the most current taxonomic status of each parasite species is noted. Interestingly, there are no data on endoparasites from more than 68% of described subterranean rodents, which indicates that discovery and documentation are at an early stage and must continue. [ABSTRACT FROM AUTHOR]

Simple Summary: The naked mole-rat is a small, long-lived mammal that lives in large colonies in subterranean burrows. The lifestyle of naked mole-rats makes their study difficult, and many details about their biology remain enigmatic. For evolutionary biologists and ecologists, a special trait exhibited by this species, their eusocial lifestyle, is of particular interest. In this study, we explored the determinants of early offspring survival using data from observations of 14 captive colonies over a total period of seven years. Our study revealed that early pup survival was significantly improved by higher pup body mass and maternal number of mammae and significantly reduced by increased maternal body mass and colony size. The latter negative effect may, however, be an artifact of the captive conditions in which colonies were kept. We further discuss the implications of the similarities and differences of naked mole-rats to eusocial insects and cooperatively breeding mammals in these determinants to shed light on the origin and maintenance of eusociality in mammals. In eusocial insects, offspring survival strongly depends on the quality and quantity of non-breeders. In contrast, the influence of social factors on offspring survival is more variable in cooperatively breeding mammals since maternal traits also play an important role. This difference between cooperative insects and mammals is generally attributed to the difference in the level of sociality. Examining offspring survival in eusocial mammals should, therefore, clarify to what extent social organization and taxonomic differences determine the relative contribution of non-breeders and maternal effects to offspring survival. Here, we present the first in-depth and long-term study on the influence of individual, maternal, social and environmental characteristics on early offspring survival in a eusocial breeding mammal, the naked mole-rat (Heterocephalus glaber). Similarly to other mammals, pup birth mass and maternal characteristics such as body mass and the number of mammae significantly affected early pup survival. In this eusocial species, the number of non-breeders had a significant influence on early pup survival, but this influence was negative—potentially an artifact of captivity. By contrasting our findings with known determinants of survival in eusocial insects we contribute to a better understanding of the origin and maintenance of eusociality in mammals. [ABSTRACT FROM AUTHOR]

Simple Summary: Good exploration and navigational skills are important for subterranean mammals because they inhabit an energetically costly environment. Damaraland mole-rats are social animals, but in captivity, they can be housed in colonies or individually. Social isolation may have effects on their exploratory behaviour and memory performance. Overall, colony-housed and individually housed mole-rats explored readily, but single queens and non-breeding females were more active, possibly related to increased anxiety-like behaviour when housed individually. All groups, except single-housed non-breeding females, also showed improvements in solving a Y-maze, while both colony and single males made fewer errors in the maze over time. Thus, learning is probable but may have been underestimated in this experiment. In general, social isolation has a limited effect on Damaraland mole-rat exploration and spatial learning. Exploratory behaviour, spatial learning and memory affect the survival of animals and appear to be dependent on the specific habitat that a species occupies. Good spatial navigation and memory are particularly important for subterranean animals, as it is energetically expensive to inhabit this niche. Damaraland mole-rats are subterranean mammals that live in colonies with organised social structures. Damaraland mole-rats have been maintained in the laboratory for many years and can be housed in groups or individually. We evaluated the effect of social isolation on the exploratory behaviour and spatial memory of single-housed and colony-housed animals and also considered potential differences in animals with different social statuses. We predicted that solitary housing would increase anxiety-like behaviour and result in higher activity and more errors when solving a maze. Exploration by colony- and single-housed mole-rats was tested in an open-field test, where all individuals explored readily. Single-housed queens and non-breeding females showed increased activity and spent more time in tunnels, which can be explained by increased anxiety. In the Y-maze, improvements in solving the maze were observed in all experimental groups, except in single-housed non-breeding females. In addition, all males showed a decrease in the number of errors in the maze. Spatial learning is thus apparent but could not be conclusively proven. It was possibly underestimated, as magnetic cues that may be used by mole-rats as stimuli for navigation were removed in the experimental setup. Overall, it appears that social isolation has a limited effect on the exploratory behaviour and spatial learning of Damaraland mole-rats. [ABSTRACT FROM AUTHOR]

Simple Summary: Naked mole-rats have a unique social system similar to insects, where there is one reproductive female, one to three reproductive males, and the rest are non-reproductive members of the colony. Within the non-reproductive members, there are dispersers that attempt to leave the colony. We aimed to investigate if naked mole-rat dispersers have consistent behavioral responses to situations and therefore possess animal personality traits. The naked mole-rat dispersers showed consistent responses to different situations, and had consistent differences between individuals. In addition, we recorded how naked mole-rats respond to situations that threaten their survival, such as a new environment and predators. Further investigation into these animal personality traits and how they compare between other colony members, such as workers and soldiers are needed to understand the naked mole-rat social system. Animal personality traits (consistent behavioral differences between individuals in their behavior across time and/or situation) affect individual fitness through facets, such as dispersal. In eusocial naked mole-rat (Heterocephalus glaber) colonies, a disperser morph may arise with distinct morphological, behavioral, and physiological characteristics. This study aimed to quantify the personality traits of a cohort of disperser morphs of naked mole-rat (NMR). Behavioral tests were performed on twelve disperser morphs (six males and six females) in an observation tunnel system that was novel and unfamiliar. Novel stimuli (fresh snakeskin, tissue paper, and conspecific of the same sex) were introduced for fifteen minutes, and the behavioral acts of the individual were recorded. A total of 30 behaviors were noted during the behavioral tests of which eight were used to quantify aggression, boldness, and exploration. The NMR disperser morphs showed consistent individual differences in boldness, and exploration across time and test, indicating a distinct personality. In addition, new naked mole-rat responses including disturbance behaviors; confront, barricade, and stay-away, were recorded. Further investigations into the relationships between animal personality traits and social hierarchy position in entire colonies are needed for more informative results as we further investigate the role of personality in cooperatively breeding societies. [ABSTRACT FROM AUTHOR]

The social mole-rats of the family Bathyergidae show elaborate social organisation that may include division of labour between breeders and non-breeders as well as across non-breeders within their groups. However, comparative behavioural data across the taxa are rare and contrasts and similarities between species are poorly understood. Field studies of social bathyergids usually involve capturing all group members until the entire group is captured. Because each animal is only captured once and traps are typically placed in close proximity to active foraging areas, the order in which animals are captured provides an indication of the foraging activity of different individuals and of the spatial organisation of the group within the burrow system. Here, we compare the association of capture order with breeding status, sex, and body mass in four species and subspecies of social bathyergids, which vary in group size and represent all three social genera within the family Bathyergidae. We show that in naked and Damaraland mole-rats (Heterocephalus glaber and Fukomys damarensis), male and female breeders are captured later than non-breeders, whereas in two different subspecies of the genus Cryptomys only female breeders are captured later than non-breeders. The effect sizes vary largely and are 10 times larger in naked mole-rats as compared to Fukomys and 3–4 times larger than in Cryptomys. Among non-breeders, sex effects are notably absent in all species and body mass predicted capture order in both naked and Damaraland mole-rats. In naked mole-rats, larger non-breeders were captured earlier than smaller ones, whereas in Damaraland mole-rats intermediate-sized non-breeders were captured first. Our data suggest that there are similarities in behavioural structure and spatial organisation across all social bathyergid species, though the most pronounced differences within groups are found in naked mole-rats.

Transmission of parasites between hosts is facilitated by close contact of hosts. Consequently, parasites have been proposed as an important constraint to the evolution of sociality accounting for its rarity. Despite the presumed costs associated with parasitism, the majority of species of African mole-rats (Family: Bathyergidae) are social. In fact, only the extremes of sociality (i.e., solitary and singular breeding) are represented in this subterranean rodent family. But how did bathyergids overcome the costs of parasitism? Parasite burden is a function of the exposure and susceptibility of a host to parasites. In this review I explore how living in sealed burrow systems and the group defenses that can be employed by closely related group members can effectively reduce the exposure and susceptibility of social bathyergids to parasites. Evidence suggests that this can be achieved largely by investment in relatively cheap and flexible behavioral rather than physiological defense mechanisms. This also shifts the selection pressure for parasites on successful transmission between group members rather than transmission between groups. In turn, this constrains the evolution of virulence and favors socially transmitted parasites (e.g., mites and lice) further reducing the costs of parasitism for social Bathyergidae. I conclude by highlighting directions for future research to evaluate the mechanisms proposed and to consider parasites as facilitators of social evolution not only in this rodent family but also other singular breeders.

Small size in mammals usually restricts long-term, frequent monitoring of endocrine function using plasma as a matrix. Thus, the non-invasive monitoring of hormone metabolite concentrations in excreta may provide an invaluable approach. The aim of the current study was to examine the suitability of enzyme immunoassays (EIAs) for monitoring responses to stressors in the naked mole-rat (Heterocephalus glaber, NMR) using urine and feces as hormone matrices. A saline control administration, and a high- and low-dose adrenocorticotropic hormone (ACTH) challenge were performed on six male and six female disperser morph NMRs. The results revealed that a 5α-pregnane-3β,11β,21-triol-20-one EIA detecting glucocorticoid metabolites (GCMs) with a 5α-3β-11β-diol structure is the most suitable assay for measuring concentrations in male urine samples, whereas an 11-oxoaetiocholanolone EIA detecting GCMs with a 5β-3α-ol-11-one structure appears the most suitable EIA for quantifying GCMs in female urine. An 11-oxoaetiocholanolone EIA detecting 11,17 dioxoandrostanes was the most suitable EIA for quantifying GCMs in the feces of both sexes. There were sex-related differences in response to the high- and low-dose ACTH challenge. We recommend using feces as a more suitable matrix for non-invasive GCM monitoring for NMRs which can be valuable when investigating housing conditions and other welfare aspects.

In eusocial insects, offspring survival strongly depends on the quality and quantity of non-breeders. In contrast, the influence of social factors on offspring survival is more variable in cooperatively breeding mammals since maternal traits also play an important role. This difference between cooperative insects and mammals is generally attributed to the difference in the level of sociality. Examining offspring survival in eusocial mammals should, therefore, clarify to what extent social organization and taxonomic differences determine the relative contribution of non-breeders and maternal effects to offspring survival. Here, we present the first in-depth and long-term study on the influence of individual, maternal, social and environmental characteristics on early offspring survival in a eusocial breeding mammal, the naked mole-rat (Heterocephalus glaber). Similarly to other mammals, pup birth mass and maternal characteristics such as body mass and the number of mammae significantly affected early pup survival. In this eusocial species, the number of non-breeders had a significant influence on early pup survival, but this influence was negative—potentially an artifact of captivity. By contrasting our findings with known determinants of survival in eusocial insects we contribute to a better understanding of the origin and maintenance of eusociality in mammals.

Exploratory behaviour, spatial learning and memory affect the survival of animals and appear to be dependent on the specific habitat that a species occupies. Good spatial navigation and memory are particularly important for subterranean animals, as it is energetically expensive to inhabit this niche. Damaraland mole-rats are subterranean mammals that live in colonies with organised social structures. Damaraland mole-rats have been maintained in the laboratory for many years and can be housed in groups or individually. We evaluated the effect of social isolation on the exploratory behaviour and spatial memory of single-housed and colony-housed animals and also considered potential differences in animals with different social statuses. We predicted that solitary housing would increase anxiety-like behaviour and result in higher activity and more errors when solving a maze. Exploration by colony- and single-housed mole-rats was tested in an open-field test, where all individuals explored readily. Single-housed queens and non-breeding females showed increased activity and spent more time in tunnels, which can be explained by increased anxiety. In the Y-maze, improvements in solving the maze were observed in all experimental groups, except in single-housed non-breeding females. In addition, all males showed a decrease in the number of errors in the maze. Spatial learning is thus apparent but could not be conclusively proven. It was possibly underestimated, as magnetic cues that may be used by mole-rats as stimuli for navigation were removed in the experimental setup. Overall, it appears that social isolation has a limited effect on the exploratory behaviour and spatial learning of Damaraland mole-rats.

Biological rhythms are rhythmic fluctuations of biological functions that occur in almost all organisms and on several time scales. These rhythms are generated endogenously and entail the coordination of physiological and behavioural processes to predictable, external environmental rhythms. The light-dark cycle is usually the most prominent environmental cue to which animals synchronise their rhythms. Biological rhythms are believed to provide an adaptive advantage to organisms. In the present review, we will examine the occurrence of circadian and seasonal rhythms in African mole-rats (family Bathyergidae). African mole-rats are strictly subterranean, they very rarely emerge aboveground and therefore, do not have regular access to environmental light. A key adaptation to their specialised habitat is a reduction in the visual system. Mole-rats exhibit both daily and seasonal rhythmicity in a range of behaviours and physiological variables, albeit to different degrees and with large variability. We review previous research on the entire circadian system of African mole-rats and discuss output rhythms in detail. Laboratory experiments imply that light remains the strongest zeitgeber for entrainment but in the absence of light, animals can entrain to ambient temperature rhythms. Field studies report that rhythmic daily and seasonal behaviour is displayed in their natural habitat. We suggest that ambient temperature and rainfall play an important role in the timing of rhythmic behaviour in mole-rats, and that they likely respond directly to these zeitgebers in the field rather than exhibit robust endogenous rhythms. In the light of climate change, these subterranean animals are buffered from the direct and immediate effects of changes in temperature and rainfall, partly because they do not have robust circadian rhythms, however, on a longer term they are vulnerable to changes in their food sources and dispersal abilities.

Females of many species discriminate among males when choosing a mate and this can bear indirect and direct benefits including the avoidance of parasite transmission from infested males. In rodents, this may be mediated by androgen hormones that affect the expression of urinary odors. Female choosiness may also vary with a female’s infestation status, with infested females being less choosy. In the current study we tested the preference of cooperatively breeding highveld mole-rat (Cryptomys hottentotus hottentotus) females for male urinary odors from healthy males and those naturally infested with a cestode (Mathevotaenia sp.). Thirty females (15 healthy, 15 infested) were allowed to explore a Y-maze with urine samples from healthy and infested males and the frequency of entering choice arms and chambers as well as the duration spend with each odor sample was recorded. Infestation status did neither affect male body mass, urinary testosterone, nor cortisol levels or the body condition of females. Although overall female activity was not affected by infestation status, infested females entered choice arms and chambers significantly less frequently than healthy females. Surprisingly, healthy females preferred odors from infested males while the opposite was true for infested females, independent of male hormone levels. As the study species lives in groups that tend to share the same infestation status, we suggest that highveld mole-rat females may exhibit a preference for unfamiliar odors, possibly as an indicator of genetic diversity, rather than discriminate between infestation status of males. Similar mechanisms may also play a role in other social species.

Although the biology of common mole-rats of the genus Fukomys has been intensively studied over the last three decades, some lineages of this speciose group of subterranean rodents remain virtually unknown to science. One of these poorly studied species is the Central African mole-rat, Fukomys ochraceocinereus (von Heuglin, 1864), which occurs in the tropical savannahs and woodlands of the northern Democratic Republic of the Congo, South Sudan, Uganda, and the Central African Republic. Here I summarize the taxonomic history and available data on the distribution and morphology of this enigmatic mole-rat, adding selected observations from museum collections. F. ochraceocinereus is a comparatively large representative of its genus that appears to express little sexual dimorphism and notable variation in pelage color. Its genetics and karyology remain severely understudied. The review highlights both apparent peculiarities of the species as well as research gaps which should be addressed by future studies on F. ochraeocinereus and its relatives, including taxonomic revisions. [ABSTRACT FROM AUTHOR]

African mole-rats (Bathyergidae) have intrigued gerontologists since extreme longevity and intraspecific divergence in ageing rates were demonstrated in some of their representatives. Although many ageing-related papers on African mole-rat species have been published in the last 20 years, a comprehensive overview of lifespan distributions, longevity metrics, and annual mortality rates in different mole-rat species (including solitary ones) is still lacking. In this paper, we aim to fill this gap by (re)analyzing published and hitherto unpublished longitudinal data of five African mole-rat species with different social organizations. We present strong indications that solitary mole-rats age faster than social species, and that the poorly studied Mashona mole-rat Fukomys darlingi exhibits some ageing characteristics not previously reported in other social mole-rats. Our study also provides the first overview of shape metrics of longevity for African mole-rat species intended to serve as a starting point for future updates. [ABSTRACT FROM AUTHOR]

In our study, we analyzed two poorly known aspects of thermal biology in subterranean rodents, the African mole-rats, that being daily energy expenditure and development of thermoregulation in juvenile mole-rat using the social Mashona mole-rat, Fukomys darlingi from southern Malawi. We performed laboratory measurements over 24 h to assess the daily energy expenditure (DEE) in adults as well as the development of the thermoregulatory abilities of juveniles at different ages. To assess the effect of ambient temperature (Ta), we exposed mole-rats to either 30 °C or 20 °C, which represents a thermoneutral as well as a thermally challenging temperature, respectively. The DEE at a Ta of 20 °C was lower than expected based on the calculation from the resting metabolic rate (RMR) at the same Ta, this suggested that heat derived from physical activity can substitute the thermoregulatory heat and thus decrease the cost on thermoregulation. To assess the development of thermoregulation in juveniles and the effect of the presence of family members on it, we measured core body temperature (Tb) and resting metabolic rate (RMR) in juveniles under several social contexts: while alone, with littermates, with littermates and mother and with littermates and both parents. Only juveniles older than one month could generate heat to keep Tb higher than Ta. Thermoregulation appeared to be fully developed in three-month-old juveniles. The presence of adult(s), but not littermates, helped to increase Tb and to decrease the RMR of juveniles. Although the results are mostly preliminary and some interpretations are limited due to low sample size in some social contexts, we may conclude that in this mole-rat species, development of thermoregulation is slow even compared to other altricial mammals. [ABSTRACT FROM AUTHOR]

In the African mole-rat family (Bathyergidae), species show both year-round and seasonal breeding patterns even though all species inhabit regions with varying seasonal rainfall patterns (a well-known selection driver of seasonal breeding). This short review suggests a novel hypothesis explaining why some African mole-rat species breed seasonally and others year-round. Namely, group-living African mole-rat species (with a mean colony size greater than five) exhibit an aseasonal reproduction pattern, as breeders, particularly females, possess year-round high body conditions. This is likely due to access to abundant and high-quality resources (food and water) throughout the year and reduced frequency of engaging in strenuous activities, such as digging, due to the increased efficiency of locating and retrieval of resource as a function of group size. While in the solitary and social species, with mean colony sizes of five or less, there are too few colony members to provide sufficient cooperative care benefits, the breeders show a seasonality to their body condition and thus breeding investment. As such, we hope to start a discussion on how group size could affect circannual breeding rhythms and propose some avenues for future research. [ABSTRACT FROM AUTHOR]

Animal personality traits (consistent behavioral differences between individuals in their behavior across time and/or situation) affect individual fitness through facets, such as dispersal. In eusocial naked mole-rat (Heterocephalus glaber) colonies, a disperser morph may arise with distinct morphological, behavioral, and physiological characteristics. This study aimed to quantify the personality traits of a cohort of disperser morphs of naked mole-rat (NMR). Behavioral tests were performed on twelve disperser morphs (six males and six females) in an observation tunnel system that was novel and unfamiliar. Novel stimuli (fresh snakeskin, tissue paper, and conspecific of the same sex) were introduced for fifteen minutes, and the behavioral acts of the individual were recorded. A total of 30 behaviors were noted during the behavioral tests of which eight were used to quantify aggression, boldness, and exploration. The NMR disperser morphs showed consistent individual differences in boldness, and exploration across time and test, indicating a distinct personality. In addition, new naked mole-rat responses including disturbance behaviors; confront, barricade, and stay-away, were recorded. Further investigations into the relationships between animal personality traits and social hierarchy position in entire colonies are needed for more informative results as we further investigate the role of personality in cooperatively breeding societies.

Rodents show a wide range of anatomical, physiological, and behavioral adaptations to life underground. Cranial and postcranial bone morphologies are deeply impacted by the modes of digging, which can involve either incisors or claws. However, the morphological variation of these elements still needs to be accurately quantified to assess the degree of specializations of the fossorial rodent families in regards to their respective evolution. Here, we focus on the morpho-functional characteristics of the masticatory apparatus in two families of subterranean rodents, the Spalacidae and Bathyergidae. We quantify skull shape in five spalacid genera using geometric morphometric methods, as well as biomechanical estimates for adductor muscles, which are compared with data previously published on bathyergids. We show that skull shape of spalacids has a greater disparity and lower biomechanical estimates than bathyergids, in which the fossorial activity, notably chisel-tooth digging, more significantly impacted the evolution of the skull. Among spalacids, Spalax shows the most extreme specializations to life underground and displays the highest number of morphological convergences with chisel-tooth digging bathyergids, especially regarding its cranial shape and high biomechanical estimate for the temporalis muscle. Fewer morphological convergences were observed between other spalacids and bathyergids. Different evolutionary histories can potentially explain discrepancies observed between the two families, the first bathyergid morphological adaptations to fossorial life being much older than those of spalacids. [ABSTRACT FROM AUTHOR]

African mole-rats of the genus Fukomys (Northern common mole-rats) combine a monogamous mating system and pronounced sexual size dimorphism; a pattern highly untypical for mammals. At the same time, they live in cooperatively breeding groups composed of reproductive and non-reproductive members of both sexes. How and to which degree sex and breeding status influence morphofunctional characters in eusocial mole-rats is not well characterized but essential to come to a comprehensive understanding of their peculiar social system. Here, we explore patterns of morphological differentiation in skulls of Ansell’s mole-rats (Fukomys anselli) by means of multivariate analysis of linear skull measurements combined with a 2D shape analysis of cranium and mandible. Compared to females, males display larger skulls relative to body size and show an expansion of the facial portion of the cranium, while reproductive status did not have an effect on any of the traits studied. We also show that species of Fukomys mole-rats display a scaling of relative sexual body size dimorphism in compliance to Rensch’s rule, which is deemed indicative of intense male intrasexual competition. For the bathyergid family as a whole, results of scaling analyses were more ambiguous, but also indicative of Rensch’s rule conformity. In line with genetic field data, our results point to a greater role of male-male conflicts in Fukomys than is traditionally assumed and support the notion that reproductive status does not correlate with morphofunctional segregation in these unusual rodents.

The relationship between muscle architectural properties and hind foot drumming of African mole-rats has yet to be determined using established methodology. Therefore, the internal structure of 32 hind limb muscles was evaluated in two drumming and one non-drumming species of Bathyergidae. The muscle mass (MM), fascicle length (Lf), and angle of pennation were measured to calculate the physiological cross-sectional area (PCSA) as well as estimate the maximum isometric force of contraction (Fmax). The most significant differences for the various muscle architecture parameters analyzed in synergistic muscle groups and individual muscles were observed between the rapid drumming Georychus capensis and the non-drumming Cryptomys hottentotus natalensis. The PCSA values of the hip extensors, hip adductors, knee extensors, and knee flexors of G. capensis were significantly larger than that of C. h. natalensis. Additionally, the hip extensors and knee flexors of both the drumming species (G. capensis and Bathyergus suillus) were shown to be capable of higher power output compared to the non-drumming species, and the hip adductors of G. capensis capable of faster contraction. M. gracilis anticus may play a key role in facilitating hind foot drumming as it was the only muscle to be significantly different in G. capensis and C. h. natalensis for all three muscle architecture parameters analyzed. Furthermore, it features in the high shortening capacity quadrant of the functional space plot of both G. capensis and B. suillus but not the non-drumming C. h. natalensis. [ABSTRACT FROM AUTHOR]

African mole‐rats (family Bathyergidae) show a wide variety of social organization, from strictly solitary to highly social. Within the family, both modes of ovulation, spontaneous and induced, have been found and are thought to relate to seasonality of reproduction and opportunities for dispersal. Damaraland mole‐rats (Fukomys damarensis) are cooperative breeders, which breed year‐round and exhibit high reproductive skew. Reproduction is monopolized by a single female (queen) in each group, and non‐reproductive females in the presence of the queen are anovulatory. The species was previously described as being a spontaneous ovulator; however, subsequent attempts to determine the length of the oestrous cycle have failed. Therefore, we revisited this subject by studying the urinary progesterone and oestradiol profiles over a period of 7 weeks and the ovarian histology of females kept under different social conditions. The first experiment shows that average urinary progesterone concentrations of breeder females paired with a vasectomized male (full physical contact) are significantly higher than those of breeder females kept singly, non‐breeder females kept singly and non‐breeder females kept within their natal group. In the second experiment, we subjected females consecutively to different experimental conditions: full physical contact (stage 1), isolation (stage 2), non‐physical contact (stage 3) and measured their steroid hormone profiles. The females showed elevated concentrations of urinary progesterone during the phase of full physical contact and basal concentrations during the subsequent single housing phase. Correspondingly, no corpora lutea of ovulation were found in the ovaries of these females in the latter phase. During stage 3, in which females were separated from a male by wire mesh (non‐physical contact), one of three females showed elevated progesterone and oestradiol concentrations throughout the sampling period and analysis of ovarian histology revealed a corpus luteum. Together, these results support the view that Damaraland mole‐rats are induced ovulators, in which non‐coital male cues are able to induce ovulation. [ABSTRACT FROM AUTHOR]

Giant mole-rats (Fukomys mechowii) are remarkably long-lived subterranean rodents (maximum recorded lifespan as reported here greater than 26 years) that live in families with one reproductive pair (breeders) and their non-reproductive offspring (non-breeders). Previous studies have shown that breeders live on average approximately twice as long as non-breeders, a finding contradicting the classic trade-off between reproduction and lifespan. Because recent evidence points to the hypothalamic-pituitary-adrenal axis as playing an important role in shaping the pace of ageing in mole-rats, we analysed the influence of the social environment of giant mole-rats on intrafamilial aggression levels, indicators of long-term stress, and, ultimately, mortality. Behavioural data indicated that family constellation, especially the presence or the absence of parents, influences agonistic behaviour. As a measure of long-term stress, we established a non-invasive method of extracting and measuring cortisol from hair of giant mole-rats. Interestingly, orphaned non-breeders exhibited significantly lower levels of cortisol and lower mortality rates than did non-breeders living with both parents. Because hypercortisolism is harmful in the long-term, intrafamilial stress could help explain the earlier onset of senescence in non-breeders, resulting in a shorter lifespan. Our findings suggest that the social environment should be considered as a further factor in ageing studies involving group-living animals. This article is part of the theme issue 'Ageing and sociality: why, when and how does sociality change ageing patterns?' [ABSTRACT FROM AUTHOR]

Naked mole‐rats express many unusual traits for such a small rodent. Their morphology, social behaviour, physiology, and ageing have been well studied over the past half‐century. Many early findings and speculations about this subterranean species persist in the literature, although some have been repeatedly questioned or refuted. While the popularity of this species as a natural‐history curiosity, and oversimplified story‐telling in science journalism, might have fuelled the perpetuation of such misconceptions, an accurate understanding of their biology is especially important for this new biomedical model organism. We review 28 of these persistent myths about naked mole‐rat sensory abilities, ecophysiology, social behaviour, development and ageing, and where possible we explain how these misunderstandings came about. [ABSTRACT FROM AUTHOR]

Sexual activity and/or reproduction are associated with a doubling of life expectancy in the long-lived rodent genus Fukomys. To investigate the molecular mechanisms underlying this phenomenon, we analyzed 636 RNA-seq samples across 15 tissues. This analysis suggests that changes in the regulation of the hypothalamic–pituitary–adrenal stress axis play a key role regarding the extended life expectancy of reproductive vs. non-reproductive mole-rats. This is substantiated by a corpus of independent evidence. In accordance with previous studies, the up- regulation of the proteasome and so-called ‘anti-aging molecules’, for example, dehydroepiandrosterone, is linked with enhanced lifespan. On the other hand, several of our results are not consistent with knowledge about aging of short-lived model organisms. For example, we found the up-regulation of the insulin-like growth factor 1/growth hormone axis and several other anabolic processes to be compatible with a considerable lifespan prolongation. These contradictions question the extent to which findings from short-lived species can be transferred to longer-lived ones. [ABSTRACT FROM AUTHOR]

Despite published reports of inter‐colony aggression and invasion in the laboratory, direct aggressive competition for resources has not previously been considered as a driving force for the large colony size in naked mole‐rats (Heterocephalus glaber). We report evidence that wild colonies of naked mole‐rats invade neighbouring colonies and thus expand their territories. Furthermore, as has been observed in laboratory colonies, invading colonies may kidnap un‐weaned pups which can subsequently be incorporated into the workforce. This is convergent with the slave‐making ants and suggests that intergroup aggression between naked mole‐rat colonies may be a strong selective pressure for group living and extreme colony size. [ABSTRACT FROM AUTHOR]

Naked mole-rats Heterocephalus glaber (NMRs) are the longest-lived rodent and also resist the normal signs of senescence. In a number of species, cellular ageing has been correlated with a reduction in telomere length, yet relatively little is known about telomeres and their age-related dynamics in NMRs and other African mole-rats. Here, we apply fluorescence in situ hybridisation (FISH) to quantify telomeric repeat sequences in the NMR, the Damaraland mole-rat, Fukomys damarensis (DMR) and the Mahali mole-rat, Cryptomys hottentotus mahali (MMR). Both terminal and non-terminal telomeric sequences were identified in chromosomes of the NMR and DMR, whilst the MMR displayed only terminal telomeric repeats. Measurements of tooth wear and eruption patterns in wild caught DMRs and MMRs, and known ages in captive bred NMRs, were used to place individuals into relative age classes and compared with a quantitative measure of telomeric fluorescence (as a proxy for telomere size). While NMRs and MMRs failed to show an age-related decline in telomeric fluorescence, the DMR had a significant decrease in fluorescence with age, suggesting a decrease in telomere size in older animals. Our results suggest that among African mole-rats there is variation between species with respect to the role of telomere shortening in ageing, and the replicative theory of cellular senescence. [ABSTRACT FROM AUTHOR]

Excavation of burrows is an extremely physically demanding activity producing a large amount of metabolic heat. Dissipation of its surplus is crucial to avoid the risk of overheating, but in subterranean mammals it is complicated due to the absence of notable body extremities and high humidity in their burrows. IR-thermography in a previous study on two species of African mole-rats revealed that body heat was dissipated mainly through the ventral body part, which is notably less furred. Here, we analyzed the dorsal and ventral skin morphology, to test if dermal characteristics could contribute to higher heat dissipation through the ventral body part. The thickness of the epidermis and dermis and the presence, extent and connectivity of fat tissue in the dermis were examined using routine histological methods, while vascular density was evaluated using fluorescent dye and confocal microscopy in the giant mole-rat Fukomys mechowii. As in other hitherto studied subterranean mammals, no subcutaneous adipose tissue was found. All examined skin characteristics were very similar for both dorsal and ventral regions: relative content of adipose tissue in the dermis (14.4 ± 3.7% dorsally and 11.0 ± 4.0% ventrally), connectivity of dermal fat (98.5 ± 2.8% and 95.5 ± 6.8%), vascular density (26.5 ± 3.3% and 22.7 ± 2.3%). Absence of large differences in measured characteristics between particular body regions indicates that the thermal windows are determined mainly by the pelage characteristics.

The Bathyergidae, commonly known as blesmols or African mole-rats, is a family of rodents well-known for their subterranean lifestyle and tunnelling behaviour. Four of the five extant bathyergid genera (Cryptomys, Fukomys, Georychus and Heliophobius) are chisel-tooth diggers, that is they dig through soil with their enlarged incisors, whereas the remaining genus (Bathyergus) is a scratch-digger, only using its forelimbs for burrowing. Heterocephalus glaber, the naked mole-rat, is also a chisel-tooth digger and was until recently included within the Bathyergidae (as the most basally branching genus), but has now been placed by some researchers into its own family, the Heterocephalidae. Given the importance of the masticatory apparatus in habitat construction in this group, knowledge and understanding of the morphology and arrangement of the jaw-closing muscles in Bathyergidae is vital for future functional analyses. Here, we use diffusible iodine-based contrast-enhanced microCT to reveal and describe the muscles of mastication in representative specimens of each genus of bathyergid mole-rat and to compare them to the previously described musculature of the naked mole-rat. In all bathyergids, as in all rodents, the masseter muscle is the most dominant component of the masticatory musculature. However, the temporalis is also a relatively large muscle, a condition normally associated with sciuromorphous rodents. Unlike their hystricomorphous relatives, the bathyergids do not show an extension of the masseter through the infraorbital foramen on to the rostrum (other than a very slight protrusion in Cryptomys and Fukomys). Thus, morphologically, bathyergids are protrogomorphous, although this is thought to be secondarily derived rather than retained from ancestral rodents. Overall, the relative proportions of the jaw-closing muscles were found to be fairly consistent between genera except in Bathyergus, which was found to have an enlarged superficial masseter and relatively smaller pterygoid muscles. It is concluded that these differences may be a reflection of the behaviour of Bathyergus which, uniquely in the family, does not use its incisors for digging.

One method burrowing animals are hypothesized to use in adapting to the presumed hypoxic subterranean environment is increasing the oxygen-carrying capacity of blood. A number of recent studies have examined hematologic parameters in laboratory-reared naked mole-rats, but not in animals living under natural atmospheric conditions. To our knowledge, blood chemistry parameters have never been systematically assessed in a fossorial mammal. In this study we examined the blood of wild naked mole-rats in Kenya and Ethiopia to determine whether their blood chemistry differs significantly from naked mole-rats born and living in captivity. We also compared our results to published values for hystricomorphs, other subterranean rodents, and surface-dwelling rodents of similar size. [ABSTRACT FROM AUTHOR]

Excavation of burrows is an extremely physically demanding activity producing a large amount of metabolic heat. Dissipation of its surplus is crucial to avoid the risk of overheating, but in subterranean mammals it is complicated due to the absence of notable body extremities and high humidity in their burrows. IR-thermography in a previous study on two species of African mole-rats revealed that body heat was dissipated mainly through the ventral body part, which is notably less furred. Here, we analyzed the dorsal and ventral skin morphology, to test if dermal characteristics could contribute to higher heat dissipation through the ventral body part. The thickness of the epidermis and dermis and the presence, extent and connectivity of fat tissue in the dermis were examined using routine histological methods, while vascular density was evaluated using fluorescent dye and confocal microscopy in the giant mole-rat Fukomys mechowii. As in other hitherto studied subterranean mammals, no subcutaneous adipose tissue was found. All examined skin characteristics were very similar for both dorsal and ventral regions: relative content of adipose tissue in the dermis (14.4 ± 3.7% dorsally and 11.0 ± 4.0% ventrally), connectivity of dermal fat (98.5 ± 2.8% and 95.5 ± 6.8%), vascular density (26.5 ± 3.3% and 22.7 ± 2.3%). Absence of large differences in measured characteristics between particular body regions indicates that the thermal windows are determined mainly by the pelage characteristics. [ABSTRACT FROM AUTHOR]

The Bathyergidae, commonly known as blesmols or African mole-rats, is a family of rodents well-known for their subterranean lifestyle and tunnelling behaviour. Four of the five extant bathyergid genera (Cryptomys, Fukomys, Georychus and Heliophobius) are chisel-tooth diggers, that is they dig through soil with their enlarged incisors, whereas the remaining genus (Bathyergus) is a scratch-digger, only using its forelimbs for burrowing. Heterocephalus glaber, the naked mole-rat, is also a chisel-tooth digger and was until recently included within the Bathyergidae (as the most basally branching genus), but has now been placed by some researchers into its own family, the Heterocephalidae. Given the importance of the masticatory apparatus in habitat construction in this group, knowledge and understanding of the morphology and arrangement of the jaw-closing muscles in Bathyergidae is vital for future functional analyses. Here, we use diffusible iodine-based contrast-enhanced microCT to reveal and describe the muscles of mastication in representative specimens of each genus of bathyergid mole-rat and to compare them to the previously described musculature of the naked mole-rat. In all bathyergids, as in all rodents, the masseter muscle is the most dominant component of the masticatory musculature. However, the temporalis is also a relatively large muscle, a condition normally associated with sciuromorphous rodents. Unlike their hystricomorphous relatives, the bathyergids do not show an extension of the masseter through the infraorbital foramen on to the rostrum (other than a very slight protrusion in Cryptomys and Fukomys). Thus, morphologically, bathyergids are protrogomorphous, although this is thought to be secondarily derived rather than retained from ancestral rodents. Overall, the relative proportions of the jaw-closing muscles were found to be fairly consistent between genera except in Bathyergus, which was found to have an enlarged superficial masseter and relatively smaller pterygoid muscles. It is concluded that these differences may be a reflection of the behaviour of Bathyergus which, uniquely in the family, does not use its incisors for digging. [ABSTRACT FROM AUTHOR]

Background We review genealogical relationships, biogeographic patterns and broad historical drivers of speciation within the Bathyergidae, a group of endemic African rodents, as well as identify key taxa which need further research. Methods We sourced comparable cytochrome b sequence data (comparable data available for all members for the Family) and geographic information for all six genera of the African subterranean rodent. This information was combined into the most comprehensive and geographically representative evolutionary study for the Bathyergidae to date. Results Species richness within the Bathyergidae appears to be underestimated, with undescribed taxa in five of the six genera. Biogeographic patterns suggest large historical distributions, which were repeatedly fragmented by major landscape changes (especially rifting, uplift and drainage evolution) since the Miocene. Aside from vicariant events, other factors (ecological specialization, population-level responses and climatic change) may have been instrumental in driving divergences in the Bathyergidae. As such, adaptive differences may exist among both populations and species across their discrete ranges, driving independent evolutionary trajectories among taxa. In addition, highly fragmented distributions of divergent (and often relict) lineages indicates the possibility of narrow endemics restricted to diminishing suitable habitats. From this, it is clear that a systematic revision of the Bathyergidae is necessary; such a revision should include comprehensive sampling of all putative taxa, the addition of genomic information to assess adaptive differences, as well as ecological information.

Ruby et al. recently analyzed historical lifespan data on more than 3200 naked mole-rats, collected over a total observation period of about 38 years (Ruby et al., 2018). They report that mortality hazards do not seem to increase across the full range of their so-far-observed lifespan, and conclude that this defiance of Gompertz's law ‘uniquely identifies the naked mole-rat as a non-aging mammal’. Here, we explain why we believe this conclusion is premature.

Seismic signalling in the form of hind foot drumming plays an integral role in the communication of several species of African mole‐rats (Bathyergidae). To produce these vibrational signals, alternating hind limbs strike the ground repetitively at high speeds by flexion and extension of the hip and knee. This descriptive study aimed to determine whether anatomical differences in hind limb osteology and/or musculature between drumming and non‐drumming species of three Bathyergidae species could be detected. Formalin‐fixed left and right hind limbs of 24 animals (N = 48) consisting of three species (n = 16 each) of two drumming species, Georychus capensis and Bathyergus suillus, and one non‐drumming species, Cryptomys hottentotus natalensis, were dissected to determine the origins and insertions of individual muscles. After dissection, all soft tissue was removed by maceration. Hind limb bones, including the pelvis, were photographed, and the exact muscle origin and insertion points were electronically mapped onto the images using imaging software. On lateral view, the acetabular position was parallel to the sacrum in G. capensis, while being more ventral in position in the other two species. The shape of the femur head was spherical and the neck defined in all species. The distal shaft of the femur was gracile and the epicondyles were robust and prominent in the non‐drumming C. h. natalensis compared with the drumming species. Shallow and relatively wide patellar grooves were observed in all three species. In the two drumming species, m. gracilis was single, whereas it was double in C. h. natalensis. In all three species, m. tensor fasciae latae was absent. The more dorsal positioning of the acetabulum in G. capensis may be needed to increase the stability of the spine and allow for more force to be exerted on the pelvis during drumming. It is unlikely that m. gracilis plays a role in drumming, as the singularity or doubling thereof is variable among rodents. It is additionally postulated that m. gluteus superficialis has taken the hip rotator role of m. tensor fasciae latae as it partially inserted onto the lateral fascia of the thigh. The more robust ilia, femoral shafts and tibiae observed in the two drumming species studied here are possible adaptations for hind foot drumming, as robust bones are able to withstand the additional biomechanical loading during drumming. [ABSTRACT FROM AUTHOR]

Background We review genealogical relationships, biogeographic patterns and broad historical drivers of speciation within the Bathyergidae, a group of endemic African rodents, as well as identify key taxa which need further research. Methods We sourced comparable cytochrome b sequence data (comparable data available for all members for the Family) and geographic information for all six genera of the African subterranean rodent. This information was combined into the most comprehensive and geographically representative evolutionary study for the Bathyergidae to date. Results Species richness within the Bathyergidae appears to be underestimated, with undescribed taxa in five of the six genera. Biogeographic patterns suggest large historical distributions, which were repeatedly fragmented by major landscape changes (especially rifting, uplift and drainage evolution) since the Miocene. Aside from vicariant events, other factors (ecological specialization, population-level responses and climatic change) may have been instrumental in driving divergences in the Bathyergidae. As such, adaptive differences may exist among both populations and species across their discrete ranges, driving independent evolutionary trajectories among taxa. In addition, highly fragmented distributions of divergent (and often relict) lineages indicates the possibility of narrow endemics restricted to diminishing suitable habitats. From this, it is clear that a systematic revision of the Bathyergidae is necessary; such a revision should include comprehensive sampling of all putative taxa, the addition of genomic information to assess adaptive differences, as well as ecological information. [ABSTRACT FROM AUTHOR]

Individual variation in growth rates often generates variation in fitness. However, the ability to draw meaningful inferences from growth data depends on the use of growth models that allow for direct comparisons of growth between the sexes, between populations, and between species. Unlike traditional sigmoid functions, a recently parameterized family of unified growth models provides a reliable basis for comparisons since each parameter affects a single curve characteristic and parameters are directly comparable across the unified family. Here, we use the unified-models approach to examine the development of sexual size dimorphism in Damaraland mole-rats (Fukomys damarensis), where breeding males are larger than breeding females. Using skeletal measurements, we show here that the larger size of male Damaraland mole-rats arises from an increased growth rate across the entire period of development, rather than through sex differences in the duration or timing of growth. Male-biased skeletal size dimorphism is not unusual among rodents, and our measures of sex differences in size in captive mole-rats are close to sexual size differences in the wild, where size dimorphism = 1.04 (male:female). We hope our study will encourage the wide use of unified growth models by mammalogists. [ABSTRACT FROM AUTHOR]

Division of labour among workers is a universal property of eusocial insect societies. For Damaraland mole-rats (Fukomys damarensis), a eusocial mammal, it was proposed that workers can be divided into morphologically distinct (in terms of body mass) subcastes of frequent and infrequent workers. Here we investigate, by using capture data from a large number of colonies of Damaraland mole-rats, if body mass is multimodally distributed, which may be indicative of worker subcastes and, further, if there is a relationship between body mass and capture order, which may be indicative of morphological specialization for colony defence. Our analysis reveals unimodal body mass distributions for both sexes. Further, there is no evidence for body mass-related differences between individuals in the capture order. These data suggest that body mass is not an indicator of behavioural specialization in Damaraland mole-rats. [ABSTRACT FROM AUTHOR]

Abstract African mole-rats (family: Bathyergidae) are strictly subterranean mammals that reside in extensive networks of underground tunnels. They are rarely, if ever, exposed to light and experience muted temperature ranges. Despite these constant conditions, the presence of a functional circadian clock capable of entraining to external light cues has been reported for a number of species. In this study, we examine a social mole-rat species, Cryptomys hottentotus mahali , to determine if it possesses a functional circadian clock that is capable of perceiving light and ambient temperature cycles, and can integrate these cues into circadian rhythms of locomotor activity and core body temperature. Eight male and eight female, non-reproductive individuals were subjected to six cycles of varying light and temperature regimes. The majority of the individuals displayed daily rhythms of locomotor activity and body temperature that are synchronised to the external light and temperature cycles. Furthermore, endogenous rhythms of both locomotor activity and core body temperature were displayed under constant conditions. Thus, we can conclude that C. h. mahali possesses a functional circadian clock that can integrate external light and temperature cues into circadian rhythms of locomotor activity and core-body temperature. Highlights • Mahali mole-rats display endogenous rhythms of locomotor activity and body temperature. • Mahali mole-rats can use both light and ambient temperature to entrain their locomotor activity rhythms. • This is the first study to use ambient temperatures ramps simulate natural temperature cycles. • There are indications that ambient temperature may be a prominent zeitgeber for circadian rhythms in this mole-rat. [ABSTRACT FROM AUTHOR]

Abstract African mole-rats are subterranean rodents, which rarely if ever leave the safety of their burrow systems. The environment of the burrows is humid, with relatively stable temperatures, and may have a hypoxic and hypercapnic atmosphere. One of crucial problems related to the subterranean way of life in mammals is avoidance of overheating, because traditional mammalian cooling mechanisms are not effective under high humidity. In African mole-rats, a variety of adaptations have evolved in response to this and other challenges of the underground ecotope. Traditionally, attention has been devoted mainly to the naked mole-rat Heterocephalus glaber , which became popular as a result of its eusociality and absence of fur, both being unique phenomena in small mammals. Despite more recent research, information on other species is still relatively limited and patchy. I review the results of studies on African mole-rats that are relevant for the understanding of their energetics and thermal biology. Attention is paid to the parameters of the burrow environment, which represent the main selection pressures shaping their physiology. In addition, an overview is given of the morphological, physiological and behavioural adaptations helping mole-rats to face temperature extremes, mechanisms by which they deal with a surplus of metabolic heat and how changes in ambient temperature influence their daily activity. The naked mole-rat is compared to its furred relatives to determine whether this species is really exceptional from the point of thermal biology. An ordination analysis was conducted using published data on mole-rat body temperature, thermoneutral zone, resting metabolic rate and thermal conductance. Most of the variability in these characteristics was found to be explained by body mass, followed by temperature characteristics of climate, but not precipitation, of the species distributional ranges. This analysis shows that the naked mole-rat is comparable to the other mole-rat species in these physiological characteristics. Highlights • African mole-rats face thermally challenging environment in their burrow systems. • Various adaptations which evolved to thrive in such conditions are reviewed here. • Body mass explains most of the variability in mole-rat energetics. • Temperature characteristics of climate have also significant influence. • More than an outlier, H. glaber is on the gradient of mole-rat thermoregulation. [ABSTRACT FROM AUTHOR]

Graphical abstract Highlights • Species diversity is underestimated in the South African bathyergid genera. • Current bathyergid clades occupy different ecological niches. • Environmental changes were instrumental in fragmenting distributions. • This study contributes to our understanding of the fossorial niche. Abstract The family Bathyergidae (comprising six genera) is a group of subterranean rodents endemic to sub-Saharan Africa. Our understanding of the evolution and species richness of the South African bathyergid genera Georychus , Bathyergus and Cryptomys is limited, with the majority of species listed as Least Concern by the IUCN Red List of Threatened Species. Genetic data suggest that several cryptic species may be present in these genera. To explore genetic and ecological distinctiveness, and evaluate taxonomic richness across the ranges of Georychus , Bathyergus and to a lesser degree, Cryptomys , as well as evaluate possible scenarios which have historically influenced evolutionary patterns, we employed four protein coding markers (one mitochondrial and three nuclear) along with distribution wide sampling schemes and large sample sizes. In addition, possible ecological differences among the different intra-generic clades were explored. Genera appear to have originated in the north-eastern interior of South Africa, following novel habitats created through the Post-African I erosion cycle and dramatic changes in climate and phytogeography. In each genus, multiple geographically discrete genetic lineages (clades) are supported by both the mitochondrial and nuclear data. These lineages bear signature of the fragmentation of wider historical distributions through major environmental changes since the middle Miocene (major uplift events, Post-African II erosion cycle, drainage evolution of major river systems, sea-level fluctuations as well as climatic changes and vegetation shifts), thereby leading to long-term isolation. Along with protracted periods of separation, it appears that ecological differences further delimit the lineages in relation to geology, phytogeographic preference, elevation, rainfall and temperature. As such, two lineages in Georychus (Clades 1 and 2) and one lineages in Cryptomys (Clade I) occur at higher elevations above the Great Escarpment (in older deposits harbouring grassland vegetation, with higher rainfall and lower daily temperatures), with the remaining lineages within these genera (Clades 3, 4 and 5 in Georychus and Clades III and IV in Cryptomys) occupying a low-land distribution with contrasting climatic and geological characteristics. Although significant differences in ecological variables were also observed between Bathyergus clades, these were not consistent, given their largely low-land distributions. Our results corroborate and expand previous suggestions that several cryptic species are present within the South African Bathyergidae. [ABSTRACT FROM AUTHOR]

Abstract A strict underground style of life is one of the greatest challenges for mammals partly due to the high energetic cost of obtaining food by digging through a mechanically resistant substrate. Any energy saving adaptation, for example the effect of social thermoregulation, is thus very important for subterranean mammals. It has also been suggested that social mammals may suffer from "isolation stress" if measured alone, because the presence of other family member(s) may decrease the stress levels and thus their metabolic rates. This phenomenon known as a socio-physiological effect should be conspicuous when the metabolism of huddling individuals is measured within a species΄ thermoneutral zone (TNZ), where no energetic costs for body warming or cooling exist. In our study, we measured the resting metabolic rates of a social species of African mole-rat, the Mashona mole-rat (Fukomys darlingi), in individuals, pairs and groups of three to eight individuals. Measurements were carried out at ambient temperature below the species' TNZ (20 °C) to test the effect of social thermoregulation and at a temperature within the species' TNZ (30 °C) to test the socio-physiological effect. In pairs, the Mashona mole-rat saved 25% of its individual energetic expenses at the temperature below the TNZ. With increasing group size, energetic savings rose up to four animals, but no savings were found in larger groups. At the temperature within the TNZ, mole-rats saved 10% of individual energetic expenses in pairs, but the difference was not significant. Also, no energetic savings were found in larger groups within the TNZ. Our results on thermoregulatory savings in the TNZ are in contrast with extremely high energetic savings found by other authors in different mole-rat species. Highlights • Energetic savings of huddling were studied in the African mole-rat Fukomys darlingi. • Mole-rats in pairs saved 25% of their energy expense due to social thermoregulation. • In larger groups the influence of social thermoregulation rose up to group of four. • There were no energetic savings at the T a within the TNZ. • No savings in the TNZ indicate the absence of a socio-physiological effect. [ABSTRACT FROM AUTHOR]

Abstract The silvery mole-rat (Heliophobius argenteocinereus) is a solitary subterranean rodent with its distribution centred mainly in miombo woodlands of eastern Africa. This part of the continent was significantly influenced by the formation of the East African Rift System (EARS) during the last 25 Mya and by pronounced climatic changes in the Plio-Pleistocene that have caused genetic differentiation leading even to speciation events in many organisms. Recently, based on analysis of mitochondrial (mt) DNA sequences, it was suggested that H. argenteocinereus is a complex of six to eight species that diverged from the early to middle Miocene. In the present study, we significantly extended the sampling, re-analysed mtDNA datasets and analysed nuclear markers with the aim to assess the evolutionary history of Heliophobius. If we do not consider the old museum samples from south-eastern Democratic Republic of Congo (very divergent short mtDNA sequences obtained from ancient DNA, requiring further study from fresh material), the genus Heliophobius is composed of three major mtDNA lineages with parapatric distribution. The Rukwa Rift (+Mbeya triple junction) at the Zambia-Tanzania border, Lake Malawi, and the Eastern Arc Mountains form the biogeographical divides among these clades. The relatively shallow differences among the mitochondrial clades, divergence dating based on the use of the fossilised birth-death ratio model and a multi-locus dataset, and a very similar pattern of genetic structure to other rodents inhabiting the same area and habitat, suggest that the evolutionary history of the extant silvery mole-rat was predominantly influenced by the climatic fluctuations in the Plio-Pleistocene. Awaiting further studies employing genomic, ecological, morphological or behavioural data, we advocate for using the single name H. argenteocinereus for all evolutionary lineages within this taxon, because (1) comparison of the genetic structure observed in mtDNA and nuclear markers suggest hybridization between at least some mtDNA lineages in the contact zones; and (2) new samples close to the type localities suggest incorrect use of previous names. [ABSTRACT FROM AUTHOR]

The endoparasite fauna of Ansell's mole-rat, Fukomys anselli (Burda, Zima, Scharff, Macholán and Kawalika) (Rodentia, Bathyergidae), was studied for the first time near Lusaka, Zambia. We recovered 7 endoparasite taxa including 5 nematodes (Hexametra sp.; Protospirura numidica Seurat, 1914; Protospirura muricola Gedoelst, 1916; Protospirura sp.; and Mammalakis zambiensis Junker, Lutermann and Mutafchiev, 2017); and 2 cestodes (Inermicapsifer sp. and Rodentolepis cf. microstoma Dujardin, 1845) from 272 individuals sampled between February 2009 and February 2010. Species richness (0–2), prevalence (14.7%), and abundance (0.44 ± 0.10) were low, probably as a result of the subterranean lifestyle of the study species. Although effect of host sex on helminth burden was largely absent, helminth burden varied with season and breeding status of the host. This may be a result of seasonal variation in burrow size in response to rainfall, but also the division of labor present in the study species. [ABSTRACT FROM AUTHOR]