Xerus Inauris Research Articles

A male Cape Ground Squirrel, Xerus inauris, grooming under the midday sun in the Kgalagadi Transfrontier Park, South Africa. Note the long whiskers (vibrissae) and the small ears, anatomical features which are very likely to be put in relation with the squirrel's burrowing habits. Photograph reproduced by permission of Emmanuel Do Linh San ‐ www.pbase.com/emmanueldolinhsan

A male Cape Ground Squirrel, <i>Xerus inauris</i>, grooming under the midday sun in the Kgalagadi Transfrontier Park, South Africa. Note the long whiskers (vibrissae) and the small ears, anatomical features which are very likely to be put in relation with the squirrel's burrowing habits. Photograph reproduced by permission of Emmanuel Do Linh San ‐ www.pbase.com/emmanueldolinhsan

Anti-snake behaviour in a facultative cooperative breeder, the Cape ground squirrel

Predator harassment is an anti-predator behaviour that may increase an individual’s risk of predation, as individuals approach, threaten and harass a potential predator, yet this behaviour is still not well understood. The Cape ground squirrel (Xerus inauris) is a highly social facultative cooperative breeder from southern Africa that harasses several species of venomous snakes. We examined whether harassment was part of alloparental care by comparing harassment behaviours among different age and sex classes in Cape ground squirrel social groups. We also assessed how individuals adjusted their behaviour dependent on levels of risk by examining the Cape ground squirrel’s harassment behaviour among non-venomous, and two species of venomous snakes. We found adult females with emerged juvenile offspring took the most risk, harassing for longer durations and at higher intensities than other group members, suggesting that snake harassment was a maternal behaviour. Females with juvenile offspring only harassed the highest risk elapid snake but increased vigilance and inspection with increasing snake risk suggesting that the Cape ground squirrel can discriminate between different types of snake predators.

Effect of mating activity and dominance rank on male masturbation among free-ranging male rhesus macaques.

The adaptive function of male masturbation is still poorly understood, despite its high prevalence in humans and other animals. In non-human primates, male masturbation is most frequent among anthropoid monkeys and apes living in multimale-multifemale groups with a promiscuous mating system. In these species, male masturbation may be a non-functional by-product of high sexual arousal or be adaptive by providing advantages in terms of sperm competition or by decreasing the risk of sexually transmitted infections. We investigated the possible functional significance of male masturbation using behavioral data collected on 21 free-ranging male rhesus macaques (Macaca mulatta) at the peak of the mating season. We found some evidence that masturbation is linked to low mating opportunities: regardless of rank, males were most likely to be observed masturbating on days in which they were not observed mating, and lower-ranking males mated less and tended to masturbate more frequently than higher-ranking males. These results echo the findings obtained for two other species of macaques, but contrast those obtained in red colobus monkeys (Procolobus badius) and Cape ground squirrels (Xerus inauris). Interestingly, however, male masturbation events ended with ejaculation in only 15% of the observed masturbation time, suggesting that new hypotheses are needed to explain masturbation in this species. More studies are needed to establish whether male masturbation is adaptive and whether it serves similar or different functions in different sexually promiscuous species.

Predator detection and dilution as benefits of associations between yellow mongooses and Cape ground squirrels

Associations among organisms are thought to form because the benefits, such as increased foraging efficiency or decreased risk of predation, outweigh any costs, such as resource competition. Though many interspecific associations have been described for closely related mammals, few studies have examined the associations between mammals in different orders. The yellow mongoose (Cynictus pencillata), a carnivore, and the Cape ground squirrel (Xerus inauris), a rodent, co-occur in arid and semi-arid South Africa where they share sleeping burrows, predators, a similar body size, and the capability to emit alarm calls in response to predators. To investigate enhanced predator avoidance as a potential benefit explaining the persistence of this association, we assessed individual mongoose vigilance alone and with squirrels or other mongooses, and with varying interspecific group size, using field observations. We also tested for responses to conspecific and heterospecific alarm calls in both study species using playback experiments. The proportion of time mongoose individuals spent vigilant decreased in the presence of squirrels or other mongooses and was negatively correlated with interspecific group size; a similar pattern was previously shown for conspecific groups of Cape ground squirrels. These results are predicted by both the dilution and collective detection hypotheses. In addition, hetero- and conspecific alarm calls elicited vigilance responses in both species. These results suggest that both species can benefit from the collective detection and dilution arising from their interspecific association and that this interspecific association could be mutualistic.

Olfactory Snake‐Predator Discrimination in the Cape Ground Squirrel

AbstractSmall mammals have a number of means to detect and avoid predators, including visual, auditory and olfactory cues. Olfactory cues are particularly important for nocturnal or fossorial species where visual cues would not be as reliable. The Cape ground squirrel (Xerus inauris) is a semi‐fossorial, diurnal mammal from southern Africa. Cape ground squirrels encounter multiple species of predatory snake that pursue individuals underground where visual and social cues are limited. We assessed whether Cape ground squirrels use odours to discriminate between snakes by presenting a non‐venomous snake, a venomous snake and a control odour collected on polyethylene cubes to 11 adult female squirrels from 11 different social groups. Cape ground squirrels responded by inspecting the cube, displaying snake harassment–associated behaviours and decreasing time spent in close proximity to snake odours when compared with a control. They also displayed discrimination between two snake species by increasing the frequency of cube inspection and increasing harassment behaviours with venomous snake odours when compared with non‐venomous snake odours. We conclude that Cape ground squirrels respond with snake‐specific antipredator behaviours when presented olfactory cues alone. Olfactory discrimination may be maintained by the decreased utility of other methods of predator detection: sight and group detection, in below‐ground encounters.

Behavioural correlates of urbanisation in the Cape ground squirrel Xerus inauris

Urbanisation critically threatens biodiversity because of habitat destruction and novel selection pressures. Some animals can respond to these challenges by modifying their behaviour, particularly anti-predator behaviour, allowing them to persist in heavily transformed urban areas. We investigated whether the anti-predator behaviour of the Cape ground squirrel Xerus inauris differed in three localities that differed in their level of urbanisation. According to the habituation hypothesis, we predicted that ground squirrels in urban areas would: (a) be less vigilant and forage more; (b) trade-off flight/vigilance in favour of foraging; and (c) have shorter flight initiation distances (FID) when approached by a human observer. Observations were made in winter and summer at each locality. As expected, ground squirrels in urbanised areas were less vigilant and had shorter FIDs but did not trade-off between foraging and vigilance. In contrast, a population in a non-urbanised locality showed greater levels of vigilance, FID and traded-off vigilance and foraging. A population in a peri-urban locality showed mixed responses. Our results indicate that Cape ground squirrels reduce their anti-predator behaviour in urban areas and demonstrate a flexible behavioural response to urbanisation.

No evidence for proteolytic venom resistance in southern African ground squirrels

Many mammalian species that interact with venomous snakes show resistances to venoms. The family Sciuridae has several North American members that harass venomous snakes and show proteolytic resistances in their sera. We examined sera collected from an African ground squirrel (Xerus inauris) against two sympatric venomous snakes (Bitis arietans and Naja annulifera) and found no support for proteolytic resistance. Our results add to our understanding of the risks in predator defense within the family Sciuridae.

Immunological sex differences in socially promiscuous African ground squirrels.

Differences in how males and females respond to foreign antigens are common across taxa. Such sexual differences in the immune system are predicted to be greater in species with high promiscuity and sociality as these factors increase the likelihood of disease transmission. Intense sperm competition is thought to further this sexual dichotomy as increased investment in spermatogenesis likely incurs additional immunological costs. Xerus inauris, a ground squirrel found throughout southern Africa, is extremely social and promiscuous with one of the highest male reproductive investments among rodents. These life-history attributes suggest males and females should demonstrate a large dichotomy in immunity. Contrary to our prediction, we found no difference in spleen mass between the sexes. However, we did find significant biases in leukocyte types and red blood cell counts, possibly reflecting responses to parasite types. Among males, we predicted greater investments in spermatogenesis would result in reduced immunological investments. We found a negative association between testes and spleen size and a positive relationship between testes and number of lice suggesting trade-offs in reproductive investment possibly due to the costs associated with spermatogenesis and immunity. We suggest when measuring sexual differences in immunity it is important to consider the effects of reproductive pressures, parasite types, and life history costs.

Seasonal Patterns of Body Temperature Daily Rhythms in Group-Living Cape Ground Squirrels Xerus inauris

Organisms respond to cyclical environmental conditions by entraining their endogenous biological rhythms. Such physiological responses are expected to be substantial for species inhabiting arid environments which incur large variations in daily and seasonal ambient temperature (Ta). We measured core body temperature (Tb) daily rhythms of Cape ground squirrels Xerus inauris inhabiting an area of Kalahari grassland for six months from the Austral winter through to the summer. Squirrels inhabited two different areas: an exposed flood plain and a nearby wooded, shady area, and occurred in different social group sizes, defined by the number of individuals that shared a sleeping burrow. Of a suite of environmental variables measured, maximal daily Ta provided the greatest explanatory power for mean Tb whereas sunrise had greatest power for Tb acrophase. There were significant changes in mean Tb and Tb acrophase over time with mean Tb increasing and Tb acrophase becoming earlier as the season progressed. Squirrels also emerged from their burrows earlier and returned to them later over the measurement period. Greater increases in Tb, sometimes in excess of 5°C, were noted during the first hour post emergence, after which Tb remained relatively constant. This is consistent with observations that squirrels entered their burrows during the day to ‘offload’ heat. In addition, greater Tb amplitude values were noted in individuals inhabiting the flood plain compared with the woodland suggesting that squirrels dealt with increased environmental variability by attempting to reduce their Ta-Tb gradient. Finally, there were significant effects of age and group size on Tb with a lower and less variable Tb in younger individuals and those from larger group sizes. These data indicate that Cape ground squirrels have a labile Tb which is sensitive to a number of abiotic and biotic factors and which enables them to be active in a harsh and variable environment.

Reproductive delay in the female Cape ground squirrel (Xerus inauris)

Abstract We studied the simultaneous effects of social and environmental factors on the age of sexual maturity to identify the major determinants of reproductive delay in the cooperatively breeding female Cape ground squirrel (Xerus inauris). We verified inhibition of sexual maturity by monitoring reproductive behaviors and hormones (using a noninvasive fecal steroid hormone analysis) in prereproductive and reproductive females from 2 populations of squirrels found in southern Africa. We found that 2 social parameters hold the most influence on the timing of female sexual maturity; specifically, the number of adult breeding female group mates and related adult male group mates were associated with a substantial inhibition of female reproductive maturity. Subadult females in social groups with 0 or 1 adult breeding female became sexually mature significantly earlier than similar-aged females in social groups with ≥2 adult breeding females. Likewise, subadult females in social groups with <2 related adult m...

Determination of an optimal dose of medetomidine-ketamine-buprenorphine for anaesthesia in the Cape ground squirrel (Xerus inauris)

The optimal dose of medetomidine-ketamine-buprenorphine was determined in 25 Cape ground squirrels (Xerus inauris) undergoing surgical implantation of a temperature logger into the abdominal cavity. At the end of anaesthesia, the squirrels were given atipamezole intramuscularly to reverse the effects of medetomidine. The mean dose of medetomidine was 67.6 +/- 9.2microg/kg, ketamine 13.6 +/- 1.9 mg/kg and buprenorphine 0.5 +/- 0.06 microg/kg. Induction time was 3.1 +/- 1.4 min. This produced surgical anaesthesia for 21 +/- 4.2 min. Atipamezole 232 +/- 92 microg/kg produced a rapid recovery. Squirrels were sternally recumbent in 3.5 +/- 2.2 min.

Vigilance and grouping in the southern African ground squirrel (Xerus inauris)

AbstractAnimals may form groups in response to the foraging–vigilance trade‐off, through enhanced predator detection (collective detection hypothesis) or reduced predation risk to the individual (dilution hypothesis), allowing individuals to decrease vigilance levels. Both hypotheses predict decreasing individual vigilance levels with increasing group size; however, the collective detection hypothesis also predicts increasing overall group vigilance with increasing group size. However, in species in which vigilance and foraging are not mutually exclusive, where vigilance may not be as costly, neither of these hypotheses may apply. Here, we examine the relationship between group size and vigilance in the social Cape ground squirrel (Xerus inauris), a species that can combine foraging and vigilance behaviours. Ten groups were observed using scan sampling, measuring both group and individual vigilance and group size. A negative relationship existed between individual vigilance and group size and a positive relationship between group vigilance and group size. Therefore, in Cape ground squirrels, vigilance seems to be costly even though it can be combined with foraging behaviours. Furthermore, group vigilance behaviour gives support to the collective detection hypothesis, whilst individual vigilance gives support to both hypotheses.

The Adaptive Function of Masturbation in a Promiscuous African Ground Squirrel

BackgroundStudies of animal mating systems increasingly emphasize female multiple mating and cryptic sexual selection, particularly sperm competition. Males under intense sperm competition may manipulate sperm quantity and quality through masturbation, which could waste sperm and decrease fertility. I examined the factors influencing masturbation by male Cape ground squirrels (Xerus inauris) in light of a number of functional hypotheses.MethodologyObservational data on a marked population of squirrels were collected in east-central Namibia using scan and all-occurrences sampling.FindingsMasturbation was far more frequent on days of female oestrus and mostly occurred after copulation. Masturbation rates were higher in dominant males, which copulate more, than in subordinates and increased with number of mates a female accepts.ConclusionsThese results suggest that masturbation in this species was not a response to sperm competition nor a sexual outlet by subordinates that did not copulate. Instead masturbation could function as a form of genital grooming. Female Cape ground squirrels mate with up to 10 males in a 3-hr oestrus, and by masturbating after copulation males could reduce the chance of infection. Sexually transmitted infections (STIs) can profoundly affect fertility, and their consequences for mating strategies need to be examined more fully.

Winter body temperature patterns in free-ranging Cape ground squirrel, Xerus inauris: no evidence for torpor

The body temperature (T(b)) of Cape ground squirrels (Xerus inauris, Sciuridae) living in their natural environment during winter has not yet been investigated. In this study we measured abdominal T(b) of eight free-ranging Cape ground squirrels over 27 consecutive days during the austral winter. Mean daily T(b) was relatively stable at 37.0 ± 0.2°C (range 33.4 to 40.2°C) despite a marked variation in globe temperature (T(g)) (range -7 to 37°C). Lactating females (n = 2) consistently had a significantly higher mean T (b) (0.7°C) than non-lactating females (n = 3) and males. There was a pronounced nychthemeral rhythm with a mean active phase T(b) of 38.1 ± 0.1°C and a mean inactive phase T(b) of 36.3 ± 0.3°C for non-lactating individuals. Mean daily amplitude of T(b) rhythm was 3.8 ± 0.2°C. T(b) during the active phase closely followed T(g) and mean active phase T(b) was significantly correlated with mean active phase T(g) (r(2) = 0.3-0.9; P < 0.01). There was no evidence for daily torpor or pronounced hypothermia during the inactive phase, and mean minimum inactive phase T(b) was 35.7 ± 0.3°C for non-lactating individuals. Several alternatives (including nocturnal huddling, an aseasonal breeding pattern and abundant winter food resources) as to why Cape ground squirrels do not employ nocturnal hypothermia are discussed.

Parasite removal increases reproductive success in a social African ground squirrel

Parasites can be detrimental to the health, longevity, and reproduction of their hosts, but these costs are rarely quantified in nature. We removed ectoparasites and endoparasites from Cape ground squirrels (Xerus inauris), a highly social sciurid in southern Africa. Parasites were removed for 3 months during the dry winter season, when fewer resources are available and the impact of parasites may be greatest. We examined changes in female body mass, reproduction, grooming, and burrow use. Female body mass did not increase with parasite removal, but reproductive success (number of offspring raised to emergence) was nearly 4 times higher for treated females. Rates of allogrooming were lower for treated females, but we were unable to detect a significant change in burrow movements, perhaps because factors other than parasites (e.g., predator avoidance) affect such movements. Gestation and lactation are the most physiologically stressful processes that females undergo, and the dramatic increase in reproductive success in treated females suggests that parasites drain resources that would otherwise be allocated to reproduction. Our results suggest that nonparasitized females put the extra energy into reproduction rather than into their own body weight, which appears to enhance juvenile survival.

Quantity versus quality: how does level of predation threat affect Cape ground squirrel vigilance?

How individuals balance time spent satisfying nutritional needs and time spent being vigilant to protect against potential predation has received abundant attention from researchers. Evidence indicates that both social conditions and predator risk affect how individuals perceive environmental threat, however, the relative influence of each type of risk remains unclear. The risk allocation hypothesis considers how individuals respond to predation risk over time by adapting their investment in high-quality or low-quality vigilance. Here we use the risk allocation hypothesis as a guide to examine how different risk factors influence vigilance quantity and quality in Cape ground squirrels, Xerus inauris. We tested how the social conditions of age, sex, distance to a safe refuge, distance from a nearest neighbour and predator risk affect the amount of time (quantity) individuals spend vigilant. Individuals were more vigilant in areas with more potential predators and when they were further from a safe refuge or nearest neighbour. We then tested whether these risk factors that affected vigilance quantity also affected vigilance quality by measuring exclusive vigilance (high-cost, high-quality) and shared-foraging–vigilance (low-cost, low-quality). Individuals in high-predation risk areas invested more time in high-cost vigilance behaviour than in low-cost vigilance/foraging behaviour compared with individuals in low-predation risk areas. Since squirrels invested in high-cost vigilance significantly more than in low-cost vigilance regardless of distance to a refuge or other squirrels, we suggest that combined foraging with vigilance compromises vigilance to lower its quality.

Hepatic capillariasis in a Cape ground squirrel (Xerus inaurus) : short communication

We report, for the first time, an incidental finding of Calodium hepaticum infestation in a sub-adult female Cape ground squirrel (Xerus inaurus). Post mortem examination of the squirrel revealed severe haemoperitoneum, splenomegaly and hepatomegaly with miliary white spots distributed diffusely throughout the hepatic parenchyma. Histologically the portal tracts in the liver showed granulomatous inflammation with fibrosis and numerous giant cells. Occasional adult worms were identified and there were multiple C. hepaticum eggs distributed diffusely throughout the portal tracts and the parenchyma. The spleen also contained C. hepaticum eggs. The genus Rattus is the primary host and reservoir of C. hepaticum, but C. hepaticum infections have been reported previously in other Sciuridae. Based on our findings, people should be cautious of the zoonotic potential of C. hepaticum, when they come into contact with the Cape ground squirrel.

The Evolution of Urgency‐Based and Functionally Referential Alarm Calls in Ground‐Dwelling Species

A major evolutionary force driving functionally referential alarm calls is the need for different strategies to escape various predator types in complex structured habitats. In contrast, a single escape strategy appears to be sufficient in less-structured open habitats, and under such conditions urgency-dependent alarm calls may be favored. Nevertheless, some species, such as meerkats (Suricata suricatta), have evolved functionally referential alarm calls despite living in open areas, using only bolt-holes for retreat. To understand the evolution of different alarm call systems, we investigated the calls of sympatric Cape ground squirrels (Xerus inauris) and compared their antipredator and foraging behavior with that of meerkats. Cape ground squirrels emitted urgency-dependent alarm calls and responded to playbacks depending on urgency, not predator type. Vigilance behavior and habitat use differed between the two species. Meerkats roam widely to find prey and for efficient foraging depend on coordinated predator vigilance and escape behavior. As herbivores with smaller territories, Cape ground squirrels depend less on coordinated antipredator behavior, and urgency-dependent alarm calls encode all essential information. We conclude that habitat complexity does not explain the evolution of functionally referential alarm calls in all species, and other constraints, such as the need to coordinate group movements to maintain foraging efficiency, could be more relevant.

The relative roles of the parasol-like tail and burrow shuttling in thermoregulation of free-ranging Cape ground squirrels, Xerus inauris

As small arid-zone mammals, Cape ground squirrels ( Xerus inauris) are unusual in being diurnally active. It is postulated that they remain active during the day by using their parasol-like tails to shade their bodies whilst foraging. However, no studies have continuously measured body temperature to determine the effect of using the tail as a parasol, relative to other thermoregulatory behaviours, such as burrow retreat. We caught four free-ranging Cape ground squirrels (673 ± 36 g) and surgically implanted miniature temperature-sensitive data loggers into their abdomens, to record body temperature every 5 min to an accuracy of 0.04 °C, before they were released back into their home range and observed for two weeks. Mean daily peak black globe temperature was 41 °C, and daily peak body temperature reached 40 °C. Ground squirrels raised their tails significantly more often at globe temperatures above 30 °C, but raising the tail did not decrease body temperature, nor prevent body temperature rising. Ground squirrels retreated to burrows, at 18 °C, significantly more often at high body temperatures and body temperature dropped 1–2 °C before re-emergence. We believe that the tail was raised to provide thermal comfort during high solar radiation exposure, and that burrow retreat was employed to dissipate a heat load and remain active diurnally.

Assessing the effects of resource availability and parity on reproduction in female Cape ground squirrels: resources do not matter

AbstractThe reproductive output of iteroparous females is influenced by parity and resource availability. To gain a better understanding of the reproductive decisions made when animals are faced with potential trade‐offs between current and future reproduction; we investigated the relationship of parity and resources availability on reproductive intervals and success by comparing the performance of primiparous and multiparous female Cape ground squirrels Xerus inauris from two study sites that differed in resource availability. Our study included the correlation of reproductive behavioral data with fecal hormonal information to establish a normative endocrine profile for pregnancy and lactation. Parity had a major influence on juvenile emergence with multiparous females having more juveniles emerge successfully. Resource availability did not influence reproductive output, either because of an interaction among multiple resource components including quantity, predictability and length of time those resources were available or because other factors (such as predation or parasitism) have a greater impact on reproductive success.