Lovers, not fighters: docility influences reproductive fitness, but not survival, in male Cape ground squirrels, Xerus inauris.

Over their lifetime, individuals may use different behavioural strategies to maximize their fitness. Some behavioural traits may be consistent among individuals over time (i.e., 'personality' traits) resulting in an individual behavioural phenotype with different associated costs and benefits. Understanding how behavioural traits are linked to lifetime fitness requires tracking individuals over their lifetime. Here, we leverage a long-term study on a multi-year living species (maximum lifespan ~ 10 years) to examine how docility (an individual's reaction to trapping and handling) may contribute to how males are able to maximize their lifetime fitness. Cape ground squirrels are burrowing mammals that live in social groups, and although males lack physical aggression and territoriality, they vary in docility. Males face high predation risk and high reproductive competition and employ either of two reproductive tactics ('natal' or 'band') which are not associated with different docility personalities. We found that although more docile individuals sired more offspring on an annual basis, docility did not affect an individual's long-term (lifetime) reproductive output. Survival was not associated with docility or body condition, but annual survival was influenced by rainfall. Our findings suggest that although docility may represent a behavioural strategy to maximize fitness by possibly playing a role in female-male associations or female mate-choice, variations in docility within our study population is likely maintained by other environmental drivers. However, individual variations in behaviours may still contribute as part of the 'tool kit' individuals use to maximize their lifetime fitness. Supplementary Information: The online version contains supplementary material available at 10.1007/s00265-023-03421-8.

The use of infrared thermography to noninvasively measure the surface temperature of polar bears during bouts of social play.

Infrared thermography or thermal imagery is a noninvasive tool that can be used to measure the temperature of surfaces. Typically, thermal imagery is used for construction or military purposes but is increasingly used as a noninvasive tool in wildlife studies. We investigated the use of thermal imagery to measure surface temperature changes as a proxy for energetic expenditure. We measured the surface temperature of polar bear (Ursus maritimus) eyes, while immobilized, to determine whether the eye is a thermal window that can accurately indicate internal temperature. We found a significant difference (2.68 ± 0.41°C) between the surface temperature of the eye measured with thermal imagery and the internal rectal temperature. Additionally, we measured surface temperature changes in polar bears after bouts of social play as a proxy for energy expenditure. Mean temperature of the eye increased by 1.34 ± 0.43°C after social play, indicating that this activity increased energy expenditure. During the fasting season, polar bears rely on fat stores, and any energy expenditure beyond what is required to travel may be costly to their survival, particularly in years of low resource availability. We conclude that thermal imagery is a useful tool to noninvasively investigate the energetics of social play.

Intraspecific variation in male mating strategies in an African ground squirrel (Xerus inauris).

Male mating strategies respond to female availability such that variation in resources that affect spatial distribution can also alter cost-benefit tradeoffs within a population. In arid-adapted species, rainfall alters reproduction, behavior, morphology, and population density such that populations differing in resource availability may also differ in successful reproductive strategies. Here, we compare two populations of Cape ground squirrels (Xerus inauris), a sub-Saharan species with year-round breeding and intense mating competition. Unlike most mammals where males resort to aggressive interactions over females, male X. inauris are tolerant of one another, relying instead on other nonaggressive pre- and postcopulatory strategies to determine reproductive success. Our findings suggest that differences in resource availability affect female distribution, which ultimately leads to intraspecific variation in male reproductive tactics and sexual morphology. Sperm competition, assessed by reproductive morphometrics, was more pronounced in our high resource site where females were distributed evenly across the landscape, whereas dominance seemed to be an important determinant of success in our low resource site where females were more aggregated. Both sites had similar mating intensities, and most males did not sire any offspring. However, our low resource site had a higher variance in fertilization success with fewer males siring multiple offspring compared with our high resource site where more individuals were successful. Our results lend support to resource models where variations in female spatial distribution attributed to environmental resources ultimately impact male reproductive behaviors and morphology.

Extreme climate event promotes phenological mismatch between sexes in hibernating ground squirrels.

Hibernating ground squirrels rely on a short active period for breeding and mass accrual, and are thus vulnerable to extreme climate events that affect key periods in their annual cycle. Here, we document how a heatwave in March 2012 led to a phenological mismatch between sexes in Richardson's ground squirrels (Urocitellus richardsonii). Females emerged from hibernation and commenced breeding earlier in 2012 relative to average female emergence. Although males had descended testes and pigmented scrota, it appeared that not all males were physiologically prepared to breed since 58.6% of males had non-motile sperm when breeding commenced. Body condition, relative testes size, and the relative size of accessory glands were significant predictors of sperm motility. Males with non-motile sperm had smaller accessory glands than males with motile sperm. There was no decrease in the number of juveniles that emerged in 2012 or female yearlings recruited in 2013, nor did juveniles emerge later than other years. The impact of this heatwave on male ground squirrels emphasizes the importance of assessing the consequences of climate change on the breeding success of hibernating species in both sexes, since the different sensitivity to external cues for emergence led to a mismatch in timing under this event.

Exploring the role of life history traits and introduction effort in understanding invasion success in mammals: a case study of Barbary ground squirrels.

Invasive species-species that have successfully overcome the barriers of transport, introduction, establishment, and spread-are a risk to biodiversity and ecosystem function. Introduction effort is one of the main factors underlying invasion success, but life history traits are also important as they influence population growth. In this contribution, we first investigated life history traits of the Barbary ground squirrel, Atlantoxerus getulus, a species with a very low introduction effort. We then studied if their invasion success was due to a very fast life history profile by comparing their life history traits to those of other successful invasive mammals. Next, we examined whether the number of founders and/or a fast life history influences the invasion success of squirrels. Barbary ground squirrels were on the fast end of the "fast-slow continuum", but their life history was not the only contributing factor to their invasion success, as the life history profile is comparable to other invasive species that do not have such a low introduction effort. We also found that neither life history traits nor the number of founders explained the invasion success of introduced squirrels in general. These results contradict the concept that introduction effort is the main factor explaining invasion success, especially in squirrels. Instead, we argue that invasion success can be influenced by multiple aspects of the new habitat or the biology of the introduced species.

Trade-offs between immunity and testosterone in male African ground squirrels.

The immunocompetence handicap hypothesis (ICHH) proposes that testosterone has both beneficial effects on male reproductive potential and negative effects by suppressing the immune system. However, support for the ICHH has been variable and an alternative hypothesis suggests that testosterone may be acting indirectly via cortisol to suppress immunity (the stress-linked ICHH). A third hypothesis is that increased energetic investment in immunity results in the suppression of testosterone. We tested these hypotheses in male Cape ground squirrels (Xerus inauris) through two separate manipulations: first, by triggering a strong immune response using a lipopolysaccharide (LPS) injection and, secondly, by increasing circulating testosterone using silastic testosterone implants. Responding to an immune challenge significantly reduced testosterone, supporting the immune suppression hypothesis, while increasing circulating testosterone had no effect on immunocompetence, body mass, ectoparasite abundances or cortisol levels, failing to support either the ICHH or stress-linked ICHH. Our results add to the increasing body of literature that challenges the ICHH, and we conclude that the trade-off between testosterone and immunity is mediated through immune activation and not through testosterone in male Cape ground squirrels. Being able to test the ICHH, stress-linked ICHH and immune suppression hypotheses in a free-ranging mammal gives us a unique opportunity to examine the mechanisms mediating this trade-off.

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.

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.

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 (T(a)). We measured core body temperature (T(b)) 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 T(a) provided the greatest explanatory power for mean T(b) whereas sunrise had greatest power for T(b) acrophase. There were significant changes in mean T(b) and T(b) acrophase over time with mean T(b) increasing and T(b) 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 T(b), sometimes in excess of 5°C, were noted during the first hour post emergence, after which T(b) remained relatively constant. This is consistent with observations that squirrels entered their burrows during the day to 'offload' heat. In addition, greater T(b) 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 T(a)-T(b) gradient. Finally, there were significant effects of age and group size on T(b) with a lower and less variable T(b) in younger individuals and those from larger group sizes. These data indicate that Cape ground squirrels have a labile T(b) which is sensitive to a number of abiotic and biotic factors and which enables them to be active in a harsh and variable environment.

Female nile hippopotamus (Hippopotamus amphibius) space use in a naturalistic exhibit.

Zoological institutions provide naturalistic exhibits for their animals in order to offer a more appealing look for visitors and give the animal the opportunity to engage in more natural behaviors. Examining space use of the animals in the naturalistic exhibit may aid in the management of these animals and inform future naturalistic exhibit design. The hippopotamus is an amphibious ungulate that spends much of its days in the wild in the water but may be found along the banks of the rivers basking in the sun. Our objective was to determine how captive female hippos utilize their exhibit by examining whether hippos selected for certain areas of a naturalistic exhibit. Scan sample data were collected on a group of nine captive female hippos housed at Disney's Animal Kingdom®. Using ArcView, the data were analyzed to determine distribution of hippos in the exhibit and their utilization of depth categories while in the water. Hippos were found to aggregate in preferred areas of the exhibit, mostly water, and selected most for water depths of 0.6-1.0 m. These results will aid in the understanding of hippopotamus space use and may aid zoological institutions in the design of naturalistic exhibits for hippos.

The adaptive function of masturbation in a promiscuous African ground squirrel.

BACKGROUND: Studies 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. METHODOLOGY: Observational data on a marked population of squirrels were collected in east-central Namibia using scan and all-occurrences sampling. FINDINGS: Masturbation 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. CONCLUSIONS: These 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.