Pangolin Thermoregulation: How Ground Pangolins Survive South Africa's Cold Winters

On a July night on the South African highveld, surface temperatures can plunge below zero. Frost blankets the grassland. Yet somewhere beneath the frozen crust, curled inside a borrowed aardvark burrow where the air holds steady at 19 degrees Celsius, a Temminck's ground pangolin sleeps through the cold with its core body temperature dipping as low as 29 degrees. It is not dying. It is conserving energy with a precision that researchers are only now beginning to understand.

A landmark 2023 study published in Conservation Physiology revealed that Temminck's pangolins are facultative heterotherms — mammals that can switch between strict body temperature regulation and a more relaxed, energy-saving mode depending on resource availability. This finding overturned the assumption that pangolins maintain a fixed internal temperature like most placental mammals, and it carries direct implications for how South Africa rehabilitates and releases the continent's most trafficked wild mammal.

A Mammal Running on Low Power

Pangolins have among the lowest basal metabolic rates of any placental mammal of comparable size. Research on captive Chinese pangolins measured average daily energy expenditure at just 52 to 61 percent of the predicted value for similarly sized eutherians. Temminck's ground pangolins, which weigh between 5 and 18 kilograms, show the same pattern. Their resting body temperature of 34 to 36 degrees Celsius sits well below the mammalian average of 37 to 38 degrees.

This low metabolic baseline is not a deficiency — it is an adaptation. As obligate myrmecophages, pangolins survive entirely on ants and termites, a protein-rich but energy-poor diet that demands an animal built for efficiency rather than speed. Every calorie matters. Where a similarly sized carnivore might burn through its energy reserves in days without food, a pangolin's slow-running metabolism extends its survival window during lean periods.

But low metabolism creates a thermal vulnerability. Generating body heat costs energy, and an animal already operating near its metabolic floor has limited capacity to increase heat production when ambient temperatures drop. This is where the pangolin's behavioural and physiological toolkit becomes critical.

The Burrow as a Thermal Refuge

Ground pangolins do not typically dig their own burrows. Instead, they occupy burrows excavated by aardvarks, springhares, and porcupines, rotating between multiple refuges across home ranges that can span 2 to 30 square kilometres depending on habitat quality. The choice of burrow is not random — it is thermal strategy.

Temperature logging at Tswalu Kalahari Reserve in the Northern Cape demonstrated that burrow interiors maintain remarkable stability. While surface temperatures at the reserve swung between 4.6 and 38.3 degrees Celsius across the year, and daily variation at the surface could exceed 20 degrees, burrow temperatures fluctuated by less than half a degree over a 24-hour cycle. Winter burrow temperatures averaged 18.96 degrees Celsius, holding 3 to 4 degrees warmer than the surface mean during the coldest months.

For an animal whose body temperature can safely dip into the high twenties, a burrow at 19 degrees is not just comfortable — it eliminates the need for active thermoregulation entirely. The pangolin can let its core temperature drift toward ambient without engaging metabolic heating, saving energy that would otherwise come from an already limited caloric budget.

Burrow depth matters. Temminck's pangolins typically use burrows that extend 1 to 5 metres below the surface, with entrance diameters of 15 to 30 centimetres. The deeper the terminal chamber, the more thermally buffered it is from surface extremes. This dependence on pre-existing burrows makes aardvark populations an indirect but critical factor in pangolin thermal survival — without aardvarks maintaining the landscape of underground refuges, pangolins lose access to their primary climate control system.

Switching from Night to Day

Temminck's ground pangolins are primarily nocturnal during the warm summer months, emerging after dark to forage when ant and termite activity peaks and predation risk from raptors is minimised. But winter changes the equation. With less prey available and colder nighttime temperatures increasing the metabolic cost of foraging, pangolins in South African habitats shift toward diurnal activity.

This seasonal reversal is not incidental. Daytime emergence exposes pangolins to solar radiation, allowing them to absorb heat passively rather than generating it metabolically. Researchers at Tswalu have observed pangolins basking in direct sunlight during winter mornings before initiating foraging bouts — behaviour functionally analogous to sun-basking in reptiles, though driven by different physiological constraints.

The shift carries trade-offs. Daytime activity increases exposure to visual predators including martial eagles, one of the few aerial threats capable of taking an adult pangolin. It also increases the risk of detection by humans, which in South Africa means potential interception by poachers or, in urban-edge habitats, encounters with domestic dogs. The fact that pangolins accept these elevated risks suggests the metabolic cost of nocturnal winter foraging is substantial enough to outweigh predation pressure.

Relaxing the Thermostat

The 2023 Conservation Physiology paper by Dr Darren Pietersen of the University of Pretoria and colleagues documented what may be the pangolin's most sophisticated cold-survival mechanism: facultative heterothermy. Using surgically implanted temperature loggers in free-ranging Temminck's pangolins at Tswalu, the team tracked core body temperatures across multiple seasons from 2015 to 2017.

During summer, when ants and termites were abundant, pangolins maintained tight homeothermic control, keeping body temperature within a narrow 34 to 36 degree band across the 24-hour cycle. During winter and dry periods, when prey was scarce, the animals relaxed this precision significantly. Body temperatures fluctuated over wider ranges, with the lowest recorded core temperature reaching 28.9 degrees Celsius in a single individual in March 2016. The highest recorded was 38.2 degrees.

This is not torpor in the classical sense — pangolins do not enter prolonged hibernation-like states. Instead, they adopt a flexible thermoregulatory strategy, maintaining just enough metabolic heat to stay functional while allowing their core temperature to track closer to ambient conditions. The energy savings are significant. Every degree of body temperature that does not need to be actively defended represents calories that can be redirected to immune function, movement, or simply extending survival until conditions improve.

Pietersen's team noted that this flexibility appears to be resource-driven rather than purely temperature-driven. Pangolins relaxed thermoregulation most during periods of low prey availability, not necessarily the coldest days. This suggests the strategy is an evolved response to the unpredictable food supply characteristic of semi-arid southern African savannahs, where ant and termite surface activity can collapse during drought or cold snaps.

Scales: Armour, Not Insulation

A common question is whether the keratin scales that cover approximately 20 percent of a pangolin's body surface provide thermal insulation. The short answer is: not meaningfully.

Pangolin scales are structurally similar to fingernails — dense, rigid plates of keratinised tissue arranged in overlapping rows. While keratin itself has modest insulating properties, the plate-like architecture of pangolin scales is optimised for mechanical protection against predator bites and ant stings, not heat retention. Unlike fur, which traps air in a low-conductivity layer against the skin, scales lie flat and create minimal dead air space.

The sparse, coarse hairs that grow between scales on the ventral surface and around the legs provide only marginal insulation. In practical terms, a pangolin's thermal envelope is closer to that of a naked mammal wearing armour plates than a fur-bearing animal. This reinforces why behavioural strategies — burrow selection, activity timing, and metabolic flexibility — are so much more important for thermal management than any passive insulation the body provides.

During hot conditions, the dense scale coverage may actually create a thermal liability by impeding evaporative cooling. Pangolins lack sweat glands on scaled surfaces and must rely on panting and heat loss through their unscaled underside. This bidirectional thermal challenge — poor insulation in cold and poor heat dissipation in heat — makes the burrow microclimate even more critical as a thermal buffer.

What This Means for Rehabilitation

South Africa's pangolin rehabilitation programmes, overseen by Prof Ray Jansen of the African Pangolin Working Group and supported by organisations including Saving the Survivors and the Johannesburg Wildlife Veterinary Hospital, handle dozens of confiscated pangolins each year. Most arrive malnourished, dehydrated, and immunocompromised from the stress of the trafficking pipeline.

For these compromised animals, thermoregulation is an immediate clinical concern. A healthy wild pangolin can afford to let its body temperature drop into the low thirties because it has fat reserves and a functioning immune system to fall back on. A trafficked pangolin with depleted energy stores and active infections cannot. Sharp temperature drops in rehabilitation enclosures can trigger pneumonia, a documented cause of mortality in captive pangolins across multiple species.

Current best practice requires maintaining enclosure temperatures between 18 and 27 degrees Celsius, with burrow-like refuges available at all times. Heat lamps or heated pads may supplement ambient temperature during cold months, but must be positioned to allow the animal to thermoregulate behaviourally — moving toward or away from the heat source as needed. Forced heating to a fixed temperature removes the animal's ability to exercise its natural heterothermic flexibility.

Release timing is equally critical. Winter releases carry elevated mortality risk because newly released pangolins must immediately establish burrow access, locate food sources, and manage thermoregulation independently, all while navigating an unfamiliar landscape. Many rehabilitation programmes now delay releases until mid-spring, when ambient temperatures exceed 15 degrees Celsius overnight and ant and termite surface activity has resumed.

The three-month soft-release protocol used at reserves like Phinda Private Game Reserve in KwaZulu-Natal includes continued supplementary feeding and monitoring during the acclimation period. Of seven pangolins released at Phinda in 2019, five survived and established home ranges — a 71 percent success rate that reflects the value of timed, supported release over abrupt translocation.

Cold Survival in a Warming World

Climate change introduces a paradox for pangolin thermoregulation. Rising average temperatures might seem beneficial for a cold-sensitive species, but the reality is more complex. Climate models project increased frequency of extreme weather events across southern Africa, including both heatwaves and unseasonal cold snaps. A pangolin adapted to predictable seasonal temperature gradients may struggle with sudden, out-of-pattern thermal stress.

More critically, changing rainfall patterns alter ant and termite availability. If prey populations crash during drought years, pangolins must extend their energy-conservation strategies for longer periods. There are physiological limits to heterothermy — a body temperature that drops too far below 28 degrees risks organ dysfunction. If the frequency and duration of prey scarcity events increase beyond what the species has historically experienced, the metabolic safety margin narrows.

Understanding pangolin thermoregulation is therefore not an academic exercise. It is foundational to predicting how the species will respond to climate change, designing effective rehabilitation protocols, and identifying which habitats will remain viable as refugia. The pangolins sleeping in their borrowed burrows beneath the frozen highveld are running a survival equation that has worked for millions of years. Whether it continues to work depends on the decisions made above ground.