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Cited by 13Physiological and biochemical adaptationsWildlife Ecology and ConservationEffects of Environmental Stressors on Livestock

Temminck’s pangolins relax the precision of body temperature regulation when resources are scarce in a semi-arid environment

Open access
Panaino, Wendy, Parrini, Francesca, Kamerman, Peter R, Hetem, Robyn S, Meyer, Leith C R, Smith, Dylan, van Dyk, Gus, Fuller, Andrea·2023·Conservation Physiology
DOI 10.1093/conphys/coad068PMC PMC10465008
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Abstract

Climate change is impacting mammals both directly (for example, through increased heat) and indirectly (for example, through altered food resources). Understanding the physiological and behavioural responses of mammals in already hot and dry environments to fluctuations in the climate and food availability allows for a better understanding of how they will cope with a rapidly changing climate. We measured the body temperature of seven Temminck’s pangolins (Smutsia temminckii) in the semi-arid Kalahari for periods of between 4 months and 2 years. Pangolins regulated body temperature within a narrow range (34–36°C) over the 24-h cycle when food (and hence water, obtained from their prey) was abundant. When food resources were scarce, body temperature was regulated less precisely, 24-h minimum body temperatures were lower and the pangolins became more diurnally active, particularly during winter when prey was least available. The shift toward diurnal activity exposed pangolins to higher environmental heat loads, resulting in higher 24-h maximum body temperatures. Biologging of body temperature to detect heterothermy, or estimating food abundance (using pitfall trapping to monitor ant and termite availability), therefore provide tools to assess the welfare of this elusive but threatened mammal. Although the physiological and behavioural responses of pangolins buffered them against food scarcity during our study, whether this flexibility will be sufficient to allow them to cope with further reductions in food availability likely with climate change is unknown.

AI snapshot

Written from the paper’s full text.

TL;DR

Temminck’s pangolins in a semi-arid environment maintained tight body temperature regulation when food was abundant but became more heterothermic when prey were scarce. They also emerged from burrows earlier during the day under low prey abundance, suggesting energy conservation prioritized over precise thermoregulation.

Key findings

  • Pangolins regulated body temperature between 34 and 36°C about 92% of the time, but showed greater 24-h body temperature variability when resources were scarce.
  • Ant abundance was 56% lower in year 1 (mean 11.4 ants/trap, SE 1.2) than year 2 (mean 20.4 ants/trap, SE 2.2); rainfall was 217 mm in year 1 vs 391 mm in year 2.
  • Lower prey abundance was associated with lower 24-h minimum body temperature at <9 ants/trap, higher 24-h maximum body temperature at <10 ants/trap, and greater 24-h body temperature amplitude at <10 ants/trap.
  • Earlier burrow emergence was associated with lower 24-h minimum body temperature, higher 24-h maximum body temperature, and lower prey abundance; the absolute minimum recorded body temperature was 28.9°C and the absolute maximum was 38.2°C.

Methods

  • Free-ranging adult Temminck’s pangolins were studied at Tswalu Kalahari Reserve, South Africa; 10 were initially located, 7 were used (4 males, 3 females, 6.5–9.4 kg), with body temperature recorded for 2938 pangolin days.
  • Core body temperature was measured with implanted temperature-sensitive data loggers recording every 5 min; activity/emergence times were obtained from camera traps and validated body temperature “notches.”
  • Prey abundance was monitored monthly using 300 pitfall traps across 30 transects, and analyses used linear mixed-effects models plus piecewise regression to identify prey abundance breakpoints.

Limitations

  • The paper states that continuous data were collected from only two animals for the full 2 years, though at least three pangolins were monitored at any given time; the small sample prevented testing for sex or body size differences.
  • The authors report small effect sizes in some models and note that body condition was not measured.
  • Weather station failure meant only 8 days of autumn 2015/2016 data were available, so autumnal data were excluded from analyses; camera traps did not always detect emergence and return events.

Model-written summary. Check it against the paper before relying on it.

Integrity signals

Registry

Crossref · 10.1093/conphys/coad068

Cites

47 of 47 references verified0 citation errors · 0 not checkable

Statistics

Nothing to recompute.

Claims the paper makes

25 of 57 cited sources checked · 14 supported · 0 flagged · accuracy not yet measured

  • Premisecitation

    Nocturnal mammals in arid zones, like Temminck's pangolin, must contend with low ambient temperatures and high radiative heat loss at night.

    “Nocturnal mammals in arid zones, like the Temminck’s pangolin, must contend with low ambient temperatures and high radiative heat loss at night (Mitchell et al., 2018; Fuller et al., 2021).”
    Quote in the paperSubstantial claim1 of 2 sources checked
    • cites Mitchell et al., 2018 source
    • cites Fuller et al., 2021 source
  • Premisecitation

    In winter, low air temperatures facilitate even greater dry heat loss.

    “In winter, low air temperatures facilitate even greater dry heat loss (Fuller et al., 2021).”
    Quote in the paperSubstantial claim1 of 1 sources checked
    • cites Fuller et al., 2021 source
  • Premisecitation

    The dry year during this study was not as extreme as had been experienced previously in the Kalahari (such as the year 2013).

    “The dry year during our study was not as extreme as had been experienced previously in the Kalahari (such as the year 2013, for example, Rey et al., 2017).”
    Quote in the paperNot a substantial claim1 of 1 sources checked
    • cites Rey et al., 2017 source
  • Premisecitation

    Food energy is required for mammals to generate the metabolic heat needed to maintain relatively high and constant body temperatures.

    “Food energy is required for mammals to generate the metabolic heat needed to maintain relatively high and constant body temperatures (Hetem et al., 2016).”
    Quote in the paperSubstantial claim1 of 1 sources checked
    • cites Hetem et al., 2016 source

      No passage in the supplied matches or retrieved text addresses whether mammals require food energy to produce metabolic heat and maintain high, constant body temperatures. Searches for those terms and related phrasing returned no substantive passage from our copy; the only result was bibliographic metadata. (LLM, conf=0.70)

      Experimental — accuracy not yet measured
  • Interpretationreasoning

    By relaxing the precision of body temperature regulation during winter, pangolins seemed to prioritize energy conservation over thermoregulation during a period when food was scarce.

    “By relaxing the precision of body temperature regulation during winter, pangolins, like many other mammals (Pereira et al., 2002; Hetem et al., 2016; Theimer et al., 2017; Weyer et al., 2020), seemed to prioritize energy conservation over thermoregulation during a period when food was scarce.”
    Quote in the paperSubstantial claim1 of 3 sources checked
    • cites Pereira et al., 2002 source
    • cites Hetem et al., 2016 source
    • cites Theimer et al., 2017 source
  • Classified as probably not a substantial claim. 38
  • May combine two claims. 38

  • Classified as probably not a substantial claim. May combine two claims.

    Premisecitation

    When food becomes less available, nocturnal mammals may respond by reducing their metabolic rate and body temperature, or by switching to diurnal activity.

    “When food becomes less available, nocturnal mammals may respond by reducing their metabolic rate and body temperature (Kelm and von Helversen, 2007; Fuller et al., 2016; Hetem et al., 2016), or by switching to diurnal activity (Van der Vinne et al., 2019; Guiden and Orrock, 2020).”
    Quote in the paperSubstantial claim2 of 3 sources checked
    • cites Kelm and von Helversen, 2007 source
    • cites Fuller et al., 2016 source

      The Physiological adjustments section says that large mammals with insufficient food availability can show lower metabolic rates and body temperatures. The Behavioural adjustments section describes some mammals shifting to nocturnal activity, but the paper does not establish this physiological response specifically for nocturnal mammals. The switch to diurnal activity is attributed to other citations in the citing sentence, not to Fuller et al. (LLM, conf=0.82)

      Experimental — accuracy not yet measured
    • cites Hetem et al., 2016 source

      No passage from our indexed copy addresses whether nocturnal mammals reduce metabolic rate and body temperature when food becomes less available. Searches for food availability, food scarcity, nocturnal activity, torpor, and metabolic rate returned no relevant passage. The diurnal-activity clause is attributed to other sources, not this paper. (LLM, conf=0.72)

      Experimental — accuracy not yet measured

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation

    Despite these responses, some aardvarks still succumbed to starvation during drought.

    “Despite these responses, some aardvark still succumbed to starvation (Rey et al., 2017; Weyer et al., 2020).”
    Quote in the paperNot a substantial claim2 of 2 sources checked
    • cites Rey et al., 2017 source

      The Discussion (§4) reports that most aardvarks died despite shifting activity patterns and having lower body temperatures. It proposes that their declining condition reflected a negative energy balance and suggests that drought reduced prey availability, but it does not establish starvation as the cause of death as definitively as the citing sentence states. (LLM, conf=0.84)

      Experimental — accuracy not yet measured
    • cites Weyer et al., 2020 source

      The Discussion reports that several aardvarks died during the 2013 drought summer and following winter despite shifts toward diurnal activity. It links those deaths to reduced prey availability, but identifies starvation as a presumed cause rather than an established one; the claim states it as fact. (LLM, conf=0.86)

      Experimental — accuracy not yet measured

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation

    During a drought in the Kalahari, aardvarks had lower 24-h minimum body temperatures compared with non-drought years.

    “During a drought in the Kalahari, aardvarks (Orycteropus afer), which also feed on ants and termites, had lower 24-h minimum body temperatures compared with non-drought years.”
    Quote in the paperNot a substantial claim1 of 1 sources checked
    • cites Weyer et al., 2020 source

  • Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.

    Premisecitation

    Switching to diurnal activity exposes mammals to higher heat loads that they would otherwise avoid, potentially leading to hyperthermia, particularly if they have insufficient body water for evaporative cooling.

    “Switching to diurnal activity also exposes mammals to higher heat loads that they would otherwise avoid, potentially leading to hyperthermia, particularly if they have insufficient body water for evaporative cooling (Weyer et al., 2020).”
    Quote in the paperNot a substantial claim1 of 1 sources checked
    • cites Weyer et al., 2020 source

      The Introduction says nocturnal mammals shifting to daytime activity face daytime heat loads and that water-deficient large mammals in hot environments exhibit higher maximum body temperatures while reducing evaporative cooling. The Discussion likewise says a shift to diurnality in hot weather increases heat exposure. Neither passage directly links that shift, combined with insufficient body water, to hyperthermia. (LLM, conf=0.82)

      Experimental — accuracy not yet measured

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation

    During a drought in the Kalahari, aardvarks shortened their active periods by a quarter.

    “They also emerged from their burrows more often during the daytime, sometimes becoming completely diurnal, and shortened their active periods by a quarter (Weyer et al., 2020).”
    Quote in the paperNot a substantial claim1 of 1 sources checked
    • cites Weyer et al., 2020 source

  • Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.

    Premisecitation

    During a drought in the Kalahari, aardvarks emerged from their burrows more often during the daytime, sometimes becoming completely diurnal.

    “They also emerged from their burrows more often during the daytime, sometimes becoming completely diurnal, and shortened their active periods by a quarter (Weyer et al., 2020).”
    Quote in the paperNot a substantial claim1 of 1 sources checked
    • cites Weyer et al., 2020 source

  • Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.

    Premisecitation

    Pietersen (2013) reported that pangolins in the Kalahari lost body condition during a winter and that some pangolins died of starvation.

    “Pietersen (2013) reported that pangolins in the Kalahari lost body condition during a winter and that some pangolins died of starvation.”
    Quote in the paperNot a substantial claim1 of 1 sources checked
    • cites Pietersen (2013)

  • Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.

    Premisecitation

    The body temperature of pangolins is lower than that of other placental mammals of similar size, likely reflecting their low metabolic rate.

    “The body temperature of pangolins has been found to be lower than that of other placental mammals of similar size, likely reflecting their low metabolic rate (McNab, 1984; Griebeler and Werner, 2016; Boyles et al., 2019).”
    Quote in the paperNot a substantial claim1 of 3 sources checked
    • cites McNab, 1984 source
    • cites Griebeler and Werner, 2016 source

      The Discussion (page unavailable) considers a possible link between body temperature and basal metabolic rate in similarly sized mammals, and the Dataset analyzed section includes Pholidota. But the paper does not report that pangolins have lower body temperatures or metabolic rates than similarly sized placental mammals; its Discussion says the temperature–metabolic-rate correlation within Pholidota was not significant. (LLM, conf=0.78)

      Experimental — accuracy not yet measured
    • cites Boyles et al., 2019 source

  • May combine two claims.

    Findingown data

    The highest absolute maximum body temperature measured in this study (38.2°C) was higher than the maximum body temperatures reported previously for any pangolin species in captivity (33.9-37.8°C) and almost 3°C higher than that previously reported for a wild Temminck's pangolin (35.4°C).

    “The highest (absolute maximum) body temperature (38.2°C) we measured was higher than the maximum body temperatures reported previously for any pangolin species in captivity (33.9–37.8°C, Jones, 1973; McNab, 1984; Heath and Hammel, 1986; Heath, 1987; Boyles et al., 2019; Challender et al., 2020; Yu et al., 2021) and almost 3°C higher than that previously reported for a wild Temminck’s pangolin (35.”
    Quote in the paperSubstantial claim1 of 3 sources checked
    • cites Jones, 1973 source

      In “Body Temperatures of Manis gigantea and Manis tricuspis” (page not available), Jones reports a maximum of 34.5°C for M. gigantea, measured while pangolins were held in cages or baskets. That supports one earlier captive measurement below 38.2°C, but this paper does not establish the full 33.9–37.8°C range or report a 35.4°C measurement for a wild Temminck’s pangolin. (LLM, conf=0.84)

      Experimental — accuracy not yet measured
    • cites McNab, 1984 source
    • cites Heath and Hammel, 1986 source

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation

    Low body temperatures may put mammals at risk of hypothermia and death, as seen in starving aardvarks.

    “Low body temperatures, however, may put mammals at risk of hypothermia and death, as seen in starving aardvarks (Rey et al., 2017; Weyer et al., 2020).”
    Quote in the paperNot a substantial claim2 of 2 sources checked
    • cites Rey et al., 2017 source
    • cites Weyer et al., 2020 source

      The Discussion (page not provided) associates lowered minimum body temperatures in aardvarks with poor body condition and high mortality, interpreting them as signs of energy deficit. The paper reports that several aardvarks died, presumably from starvation, but does not establish that low body temperature caused hypothermia or death. (LLM, conf=0.86)

      Experimental — accuracy not yet measured

  • Classified as probably not a substantial claim. May combine two claims.

    Premisecitation

    The amplitude of the 24-h rhythm of body temperature has been proposed as a proxy for the physiological welfare of mammals, with many mammals exhibiting homeothermy only when they have sufficient food and water and are not energetically challenged.

    “The amplitude of the 24-h rhythm of body temperature has been proposed as a proxy for the physiological welfare of mammals (Hetem et al., 2016; Maloney et al., 2017; Maloney et al., 2019), with many mammals exhibiting homeothermy only when they have sufficient food and water and are not energetically challenged.”
    Quote in the paperSubstantial claim2 of 3 sources checked
    • cites Hetem et al., 2016 source

      Split into 2 atoms — 0 confirmed, 1 partially supported, 0 disputed, 0 misattributed, 1 unverifiable. Expand the claim to see per-atom verdicts.

      Experimental — accuracy not yet measured
    • cites Maloney et al., 2017 source
    • cites Maloney et al., 2019 source

  • May combine two claims.

    Findingown data

    Pangolins in this study had mean body temperatures higher than those measured previously for Temminck's pangolins in captivity (32-34°C) and for one animal in the wild (32-35°C).

    “Pangolins in our study, however, had mean body temperatures that were higher compared with those measured previously for Temminck’s pangolins in captivity (32–34°C; Boyles et al., 2019; Wicker et al., 2020) and for one animal in the wild (32–35°C; Pietersen, 2013).”
    Quote in the paperSubstantial claim1 of 3 sources checked
    • cites Boyles et al., 2019 source
    • cites Wicker et al., 2020 source
    • cites Pietersen, 2013

  • May combine two claims.

    Findingcitation

    The lowest body temperature recorded for a pangolin in this study (28.9°C in March 2016) was within the range of minimum body temperatures recorded previously for captive animals (~27.6-33.4°C) and similar to that recorded for a wild pangolin (29.5°C).

    “The lowest body temperature recorded for a pangolin in our study (28.9°C in March 2016) was within the range of minimum body temperatures recorded previously for captive animals (~27.6–33.4°C; McNab, 1984; Heath and Hammel, 1986; Heath, 1987; Yu et al., 2021) and similar to that recorded for a wild pangolin (29.5°C; Pietersen, 2013).”
    Quote in the paperNot a substantial claim1 of 3 sources checked
    • cites McNab, 1984 source
    • cites Heath and Hammel, 1986 source
    • cites Pietersen, 2013

  • Classified as probably not a substantial claim. May combine two claims.

    Premisecitation

    Although food energy is important for the maintenance of high body temperatures, sufficient body water is needed for evaporative cooling.

    “Although food energy is important for the maintenance of high body temperatures, sufficient body water is needed for evaporative cooling (Ostrowski et al., 2003; Fuller et al., 2016; Hetem et al., 2016).”
    Quote in the paperSubstantial claim2 of 3 sources checked
    • cites Ostrowski et al., 2003 source
    • cites Fuller et al., 2016 source
    • cites Hetem et al., 2016 source

      No passage from the paper’s body was returned for searches on food energy, water availability, and evaporative cooling. The indexed results contain only abstract and bibliographic material, which cannot establish whether the paper supports this claim under the evidence rule. (LLM, conf=0.72)

      Experimental — accuracy not yet measured

  • May combine two claims.

    Premisecitation

    Similar findings were reported for aardvarks in the Kalahari; when drought conditions reduced grass, the availability of termites (Hodotermes mossambicus) decreased, and aardvarks exhibited greater amplitudes of the 24-h rhythm of body temperature.

    “Similar findings were reported for aardvarks in the Kalahari; when drought conditions reduced grass, the availability of termites (Hodotermes mossambicus) decreased (Weyer, 2018), and aardvarks, which also obtain water from food, exhibited greater amplitudes of the 24-h rhythm of body temperature (Weyer et al., 2020).”
    Quote in the paperNot a substantial claim1 of 2 sources checked
    • cites Weyer, 2018 source
    • cites Weyer et al., 2020 source

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Findingown data

    Small effect sizes were detected in some of the models.

    “We also detected small effect sizes in some of our models.”
    Quote in the paperNot a substantial claim

  • May combine two claims.

    Findingown data

    Pangolins maintained homeothermy (between 34 and 36°C) throughout most of the year, but exhibited heterothermy during winter.

    “Pangolins maintained homeothermy (between 34 and 36°C) throughout most of the year, but they exhibited heterothermy during winter.”
    Quote in the paperSubstantial claim

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Findingown data

    On one occasion, a pangolin was observed drinking from a cup-like woody structure at the base of a shrub that had filled with water after a rain event during summer, suggesting that they will opportunistically drink water at times.

    “On one occasion, we observed a pangolin drinking from a cup-like woody structure at the base of a shrub that had filled with water after a rain event during summer, suggesting that they will opportunistically drink water at times.”
    Quote in the paperNot a substantial claim

  • May combine two claims.

    Findingown data

    The maximum amplitude of the 24-h rhythm of body temperature was 6.8°C, with 24-h body temperature increasing from a minimum of 30.2°C to a maximum of 37.0°C within 2 h and 39 min during July 2016.

    “The maximum amplitude of the 24-h rhythm of body temperature was 6.8°C, with 24-h body temperature increasing from a minimum of 30.2°C to a maximum of 37.0°C (within 2 h and 39 min) during July 2016.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Findingreasoning

    Pangolins shifted from a predominantly nocturnal pattern during summer to greater diurnal activity during winter, presumably to reduce the metabolic costs of staying warm at night.

    “Pangolins shifted from a predominantly nocturnal pattern during summer to greater diurnal activity during winter, presumably to reduce the metabolic costs of staying warm at night.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Findingown data

    Ant abundance was lowest during autumn and winter, and highest during summer and spring.

    “Averaging across seasons, ant abundance was 56% lower during year 1 (mean: 11.4 ants/trap, SE: 1.2) compared with year 2 (mean: 20.4 ants/trap, SE: 2.2), and was lowest during autumn (mean: 9.9 ants/trap, SE: 1.3) and winter (mean: 8.2 ants/trap, SE: 1.1), and highest during summer (mean: 20.2 ants/trap, SE: 3.1) and spring (mean: 26.9 ants/trap, SE: 3.4).”
    Quote in the paperNot a substantial claim

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Findingown data

    When averaged across years, 24-h mean, minimum and maximum black globe temperatures were greatest in summer and lowest in winter.

    “When averaged across years, 24-h mean, minimum and maximum black globe temperatures were greatest in summer and lowest in winter (Supplementary Material, Tables S4, S6, and S10).”
    Quote in the paperNot a substantial claim

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Findingown data

    Greater rainfall was recorded in year 2 (391 mm) compared with year 1 (217 mm).

    “Greater rainfall was recorded in year 2 (391 mm) compared with year 1 (217 mm) (Supplementary Material, Table S12, Fig. S12).”
    Quote in the paperNot a substantial claim

  • May combine two claims.

    Findingown data

    Pangolins in this study had 24-h mean body temperatures as low as 32°C in the year with food scarcity, ~1°C lower than in the year with available food.

    “Pangolins in our study had 24-h mean body temperatures as low as 32°C in the year with food scarcity, ~1°C lower than in the year with available food.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Findingown data

    During winter, and during the year with less rainfall, which coincided with less abundant prey resources, pangolins became more heterothermic, as indicated by decreased 24-h body temperature minima and increased 24-h body temperature maxima.

    “During winter, and during the year with less rainfall, which coincided with less abundant prey resources, they became more heterothermic, as indicated by decreased 24-h body temperature minima and increased 24-h body temperature maxima.”
    Quote in the paperSubstantial claim

  • Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.

    Findingown data

    Across the 2-year period, median rainfall was greatest in summer (37 mm [8-76]) and autumn (26 mm [12-33]) months, and lowest in spring (5 mm [0-12]) and winter months (0 mm [0-0]).

    “Across the 2-year period, median (interquartile range [IQR]) rainfall was greatest in summer (37 mm [8–76]) and autumn (26 mm [12–33]) months, and lowest in spring (5 mm [0–12]) and winter months (0 mm [0–0]) (Supplementary Material, Table S11, Fig. S10, Fig. S11).”
    Quote in the paperNot a substantial claim

  • May combine two claims.

    Findingown data

    Seasonally, when averaged across years, summer was the season when pangolins had the highest 24-h minimum body temperature (mean: 33.7°C) and winter was when body temperature was lowest (mean: 32.9°C).

    “Seasonally, when averaged across years, summer was the season when pangolins had the highest 24-h minimum body temperature (mean: 33.7°C, SE: 0.17) and winter was when body temperature was lowest (mean: 32.9°C, SE: 0.17) (Supplementary Material, Tables S15 and S16, Fig. S15).”
    Quote in the paperNot a substantial claim

  • Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.

    Findingown data

    Black globe temperatures ranged from a minimum of −5.1°C in winter to a maximum of 49.5°C in summer in year 1, and from −4.2°C in winter to 50.4°C in summer in year 2.

    “Black globe temperatures ranged from a minimum of −5.1°C (winter) to a maximum of 49.5°C (summer) in year 1, and from a minimum of −4.2°C (winter) to a maximum of 50.4°C (summer) in year 2.”
    Quote in the paperNot a substantial claim

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Findingown data

    Although all these associations were statistically significant, the relationships described by the model were consistent with small effect sizes.

    “Although all these associations were statistically significant, the relationships described by the model were consistent with small effect sizes.”
    Quote in the paperNot a substantial claim

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Findingown data

    When averaged across seasons, the 24-h mean black globe temperature in year 2 was 2.1°C greater compared with year 1.

    “When averaged across seasons, the 24-h mean black globe temperature in year 2 was 2.1°C (SE: 0.3) greater compared with year 1 (Supplementary Material, Tables S9 and S10, Fig. S9).”
    Quote in the paperNot a substantial claim

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Findingown data

    Daily amplitude of black globe temperatures did not change across the seasons.

    “Daily amplitude of black globe temperatures did not, however, change across the seasons (Supplementary Material, Table S8).”
    Quote in the paperNot a substantial claim

  • May combine two claims.

    Findingown data

    The absolute minimum 24-h body temperature recorded was 28.9°C (March 2016) and the absolute maximum 24-h body temperature recorded was 38.2°C (September 2017).

    “The absolute minimum 24-h body temperature recorded was 28.9°C (March 2016) and the absolute maximum 24-h body temperature recorded was 38.2°C (September 2017).”
    Quote in the paperNot a substantial claim

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Findingown data

    The mean difference for 24-h maximum black globe temperatures between year 1 and year 2 was 0.7°C.

    “The mean difference for 24-h maximum black globe temperatures between year 1 and year 2 was 0.7°C (SE: 0.4) (Supplementary Material, Tables S5 and S6, Fig. S7).”
    Quote in the paperNot a substantial claim

  • May combine two claims.

    Findingnone given

    Pangolins regulated core body temperature within a narrow 24-h range when food resources were abundant but relaxed the precision of body temperature regulation when food resources became scarcer.

    “In conclusion, pangolins regulated core body temperature within a narrow 24-h range when food resources were abundant but relaxed the precision of body temperature regulation when food resources became scarcer.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Findingown data

    At the study site, water holes were present, but in 7 years of field work studying pangolins, the authors never observed one drinking from these sources.

    “At our study site, water holes were present, but in 7 years of being in the field there studying pangolins for various purposes, we have never observed one drinking from these sources.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Premisecitation

    Maintenance of a relatively high body temperature at night requires a high metabolic rate, which in turn requires sufficient food energy.

    “Maintenance of a relatively high body temperature at night requires a high metabolic rate, which in turn requires sufficient food energy (Jessen, 2001; Silva, 2005).”
    • Also stated in: § Abstract · plain
      “Pangolins need energy from food to maintain a constant and high body temperature, especially as they normally are active on cold nights.”
    Quote in the paperSubstantial claim0 of 2 sources checked
    • cites Jessen, 2001 source
    • cites Silva, 2005 source

  • May combine two claims.

    Interpretationreasoning

    Pangolins exhibited rapid increases in body temperature during winter when prey abundance was low (<10 ants/trap) and when they emerged before nightfall, suggesting that they may have had insufficient body water to prevent the rise in body temperature.

    “Indeed, pangolins exhibited rapid increases in body temperature during winter when prey abundance was low (<10 ants/trap) and when they emerged before nightfall, suggesting that they may have had insufficient body water to prevent the rise in body temperature.”
    Quote in the paperSubstantial claim

  • Classified as probably not a substantial claim. May combine two claims.

    Interpretationown data

    Body temperature measurement using biologging, and insect abundance using pitfall trapping may be used as tools for understanding the physiological welfare of pangolins.

    “We have shown that body temperature measurement using biologging, and insect abundance using pitfall trapping may be used as tools for understanding the physiological welfare of pangolins.”
    Quote in the paperSubstantial claim

  • May combine two claims.

    Premisecitation

    Pangolins lack UCP1 and therefore cannot implement non-shivering thermogenesis.

    “Pangolins also lack UCP1 (Gaudry et al., 2018) and therefore cannot implement non-shivering thermogenesis.”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites Gaudry et al., 2018 source

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation

    An increased breathing rate was observed in Chinese pangolins (Manis pentadactyla) when air temperatures rose >32°C.

    “an increased breathing rate was observed in Chinese pangolins (Manis pentadactyla) when air temperatures rose >32°C (Heath and Hammel, 1986).”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites Heath and Hammel, 1986 source

  • May combine two claims.

    Study design and conductcitation

    Ants that fell into pitfall traps were used as a proxy for pangolin prey abundance because ants dominated the pangolin diet and termites rarely fell into the traps.

    “Because ants dominated the pangolin diet (Panaino et al., 2022) and termites rarely fell into the traps, only the ants that fell into the traps were counted and used as a proxy for pangolin prey abundance.”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Panaino et al., 2022 source

  • Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.

    Premisecitation

    Temminck's pangolins occur in the same environment as aardvarks and have a similar diet.

    “Temminck’s pangolins occur in the same environment as aardvarks and have a similar diet (Weyer, 2018; Panaino et al., 2022), but their shallower foraging habits may make them vulnerable to fluctuations in food availability.”
    Quote in the paperNot a substantial claim0 of 2 sources checked
    • cites Weyer, 2018 source
    • cites Panaino et al., 2022 source

  • Classified as probably not a substantial claim.

    Premisecitation

    Evaporative cooling is the only way to lose heat when operative temperature exceeds an animal's surface temperature.

    “Evaporative cooling is the only way to lose heat when operative temperature exceeds an animal’s surface temperature (Mitchell et al., 2018).”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Mitchell et al., 2018 source

  • May combine two claims.

    Premisecitation

    Pangolins rarely actively seek freestanding water and obtain almost all their body water through their diet.

    “Pangolins rarely actively seek freestanding water and obtain almost all their body water through their diet (Pietersen et al., 2020).”
    Quote in the paperSubstantial claim0 of 1 sources checked
    • cites Pietersen et al., 2020 source

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation

    Pangolins lack sweat glands.

    “Pangolins lack sweat glands (Hyman and Wake, 1992)”
    Quote in the paperNot a substantial claim0 of 1 sources checked
    • cites Hyman and Wake, 1992

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Study design and conductown data

    Body temperature was recorded for seven pangolins for a total of 2938 pangolin days across the study period.

    “Body temperature was recorded for seven pangolins for a total of 2938 pangolin days across the study period (November 2015 to October 2017; Fig. 1).”
    Quote in the paperNot a substantial claim

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Study design and conductown data

    Linear mixed-effects models were assessed for linearity, heteroskedasticity, outliers, normality of residuals and multicollinearity.

    “All models were assessed for linearity (residual vs fitted plot), heteroskedasticity (residual vs fitted plot), outliers (Cook D > 1), normality of residuals (QQ-plot) and multicollinearity (variance inflation factor >5) (R package: ‘performance’, Lüdecke et al., 2021).”
    Quote in the paperNot a substantial claim

  • Probably not a substantial claim: may not pass S2 (checkable), S4 (based). May combine two claims.

    Interpretationnone given

    Body temperature biologging is invasive and difficult to implement widely.

    “Body temperature biologging, however, is invasive and difficult to implement widely.”
    Quote in the paperNot a substantial claim

  • Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.

    Study design and conductown data

    The data loggers drifted in time by ±1 min per month; therefore, appropriate time shift corrections were made before data analysis.

    “The data loggers drifted in time by ±1 min per month; therefore, appropriate time shift corrections were made before data analysis.”
    Quote in the paperNot a substantial claim

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Study design and conductown data

    Data were collected continuously from only two animals for the full 2 years because contact was lost with some pangolins.

    “we were able to collect data continuously from only two animals for the full 2 years because we lost contact with some pangolins.”
    Quote in the paperNot a substantial claim

  • May combine two claims.

    Interpretationreasoning

    Lower 24-h minimum body temperatures reduce the temperature difference between the animal's body surface and the environment, thereby reducing dry heat loss and the energy demand for heat production.

    “Another advantage of the resulting lower 24-h minimum body temperatures is that the temperature difference between the animal’s body surface and the environment would be lower, thereby reducing dry heat loss to the environment, and reducing the energy demand for heat production.”
    Quote in the paperSubstantial claim

  • Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.

    Premisecitation

    At the same time, the aardvarks' body condition deteriorated, and some succumbed.

    “At the same time, the aardvarks’ body condition deteriorated, and some succumbed.”
    Quote in the paperNot a substantial claim

  • Classified as probably not a substantial claim.

    Study design and conductnone given

    The waxed data loggers were calibrated before implantation and after explantation against a highly accurate thermometer in a water bath over a range of 26-42°C.

    “We calibrated the waxed data loggers (mass ± 20 g) before implantation and after being explanted from animals (see below) against a highly accurate thermometer (Quat 100, Heraeus, Germany) in a water bath over a range of 26–42°C.”
    Quote in the paperSubstantial claim

  • Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.

    Study design and conductown data

    Contact was lost with three of ten tagged pangolins, so seven pangolins were used in the study.

    “Contact was lost with three pangolins (one male and two females) subsequent to release. Therefore, seven pangolins (four males and three females) were used in our study.”
    Quote in the paperNot a substantial claim

  • May combine two claims.

    Interpretationreasoning

    Because pangolins get most of their body water from their food, when they experienced energy deficits as a consequence of reduced prey abundance, they likely also experienced water deficits.

    “Because pangolins get most of their body water from their food, when they experienced energy deficits as a consequence of reduced prey abundance, they likely also experienced water deficits.”
    Quote in the paperSubstantial claim
    • Rests on an assumption: Prey abundance is a proxy for food energy and water availability.

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Study design and conductcitation

    Where heteroskedasticity was detected, robust standard errors were calculated.

    “Where heteroskedasticity was detected, robust standard errors were calculated (R packages: ‘parameters’ and ‘sandwich’, Lüdecke et al., 2020, Zeileis, 2006 and Zeileis et al., 2020).”
    Quote in the paperNot a substantial claim

  • Probably not a substantial claim: may not pass S2 (checkable), S3 (load-bearing).

    Interpretationreasoning

    Biologging of body temperature is particularly useful for assessing the physiological welfare of pangolins, given that pangolins are difficult to monitor because they occur at low densities, are nocturnal, usually solitary and fossorial.

    “Biologging of body temperature is particularly useful for assessing the physiological welfare of pangolins, given that pangolins are difficult to monitor because they occur at low densities, are nocturnal, usually solitary and fossorial.”
    Quote in the paperNot a substantial claim

  • Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.

    Study design and conductreasoning

    Pangolin body condition was not measured in this study and could not be visually observed given the scale covering on the animals.

    “Although pangolin body condition was not measured in our study and could not be visually observed given the scale covering on the animals”
    Quote in the paperNot a substantial claim

  • May combine two claims.

    Study design and conductreasoning

    Camera traps did not always detect time of emergence and return because they did not always trigger and pangolins did not always return to the same burrow.

    “The camera traps did not always trigger for emergence and return events, and pangolins did not always return to the same burrow, thus the camera traps did not always detect time of emergence and return.”
    Quote in the paperNot a substantial claim

  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Interpretationreasoning

    The differences between captive and wild pangolins may explain the higher maximum body temperature recorded in this study.

    “which may explain the higher maximum body temperature recorded in our study.”
    Quote in the paperNot a substantial claim
    • Rests on an assumption: Captive pangolins are not as active as wild pangolins.
    • Rests on an assumption: Captive pangolins may be in poor condition.
    • Rests on an assumption: Captive pangolins likely are not exposed to high radiant heat loads like the pangolins in this study were when they emerged from their burrows diurnally.
60 of 60

  • The authors state this as an assumption.

    Little is known about how Temminck's pangolins respond to changing environments, and how they regulate body temperature and their activity patterns.

    “Little is known about how Temminck’s pangolins respond to changing environments, and how they regulate body temperature and their activity patterns.”

  • The paper relies on this without support.

    Temminck's pangolins' shallower foraging habits may make them vulnerable to fluctuations in food availability.

    “Temminck’s pangolins occur in the same environment as aardvarks and have a similar diet (Weyer, 2018; Panaino et al., 2022), but their shallower foraging habits may make them vulnerable to fluctuations in food availability.”

  • The authors state this as an assumption.

    Prey abundance was consistent for the month.

    “with the assumption that prey abundance was consistent for the month”

  • The paper relies on this without support.

    Captive pangolins are not as active as wild pangolins.

    “Pangolins housed under captive conditions are not as active as wild pangolins”

  • The paper relies on this without support.

    Captive pangolins may be in poor condition.

    “may be in poor condition”

  • The paper relies on this without support.

    Captive pangolins likely are not exposed to high radiant heat loads like the pangolins in this study were when they emerged from their burrows diurnally.

    “likely are not exposed to high radiant heat loads like the pangolins in our study were when they emerged from their burrows diurnally”

  • The paper relies on this without support.

    Even during winter, black globe temperatures can be as high as 37°C in the late afternoon in the Kalahari.

    “Even during winter, black globe temperatures can be as high as 37°C in the late afternoon in the Kalahari.”

  • The paper relies on this without support.

    Climate change in drylands likely is an additional threat to the survival of these threatened mammals.

    “climate change in drylands likely is an additional threat to the survival of these threatened mammals.”

  • The paper relies on this without support.

    This study is the first to investigate the body temperature patterns of wild pangolins.

    “Although our study is the first to investigate the body temperature patterns of wild pangolins”

  • The paper relies on this without support.

    Prey abundance is a proxy for food energy and water availability.

    “prey abundance, and hence food energy and water, was low”

  • The paper relies on this without support.

    Pangolins lack sufficient food energy when prey abundance is low.

    “given that the pangolins lack sufficient food energy”
11 of 11

Self-check

Percentages, sums and reported differences re-computed from the paper’s own numbers.

Each contradiction between two passages listed here was confirmed by an independent check.

  • Year 2 body-temperature amplitude has the wrong direction

    Major

    The reported year 2 minimum is 0.35°C higher and its maximum is 0.27°C lower. Those changes imply a smaller, not greater, maximum-minus-minimum amplitude in year 2.

  • Table 2 intercept t-statistic does not match its coefficient and SE

    Minor

    A coefficient of 36.18 divided by its reported SE of 0.18 gives a t-statistic of 201, not 112.28; rounding of the coefficient and SE cannot account for the difference.

    Reported 112.28 · recomputed 201

  • Discussion reverses the prey-maximum-temperature relationship

    Minor

    The discussion says lower prey abundance was associated with a lower 24-h maximum body temperature, whereas the Results say maximum body temperature increased as prey abundance decreased below 10 ants per trap.

  • Table 2 gives a positive t-statistic for a negative prey coefficient

    Minor

    A coefficient with a negative confidence interval must have a negative coefficient-to-SE t-statistic, but the table reports +14.51.

4 of 4

Cites

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Related

Authors
Panaino, W., Parrini, F., Kamerman, P. R., Hetem, R. S., Meyer, L. C. R., Smith, D., van Dyk, G., & Fuller, A.
Affiliations
  • University of the Witwatersrand (ZA)
  • Tswalu Kalahari Reserve , van Zylsrus, 8467, Northern Cape, South Africa
Corresponding author
Panaino, Wendy
ORCID
Venue
Conservation Physiology
Publisher
Oxford University Press (OUP)
Published
2023-01-01
Volume
11
Issue
1
Pages
coad068-coad068
ISSN
2051-1434 (ISSN-L)
DOI
10.1093/conphys/coad068
PMC
PMC10465008
OpenAlex
W4386261365
Semantic Scholar
4eb74fe6b6d38212ca7744d00e340149096c9bd8
Cited by
12
References
46
Citations by year
2026: 7 · 2025: 2 · 2024: 2 · 2023: 1
FWCI
2.10
Type
journal article
Language
en
Open access
Licence
Locations
  • Conservation Physiology (publishedVersion, cc-by)
  • PubMed Central (publishedVersion, cc-by)
  • PubMed (publishedVersion)
  • Massey Research Online (Massey University) (submittedVersion, cc-by)
  • UpSpace Institutional Repository (University of Pretoria) (submittedVersion, cc-by)
Funders
  • Brain Function Research Group, University of the Witwatersrand
  • Tswalu Foundation; Save Pangolins
Topics
  • Physical Sciences › Environmental Science › Ecology › Physiological and biochemical adaptations
  • Physical Sciences › Environmental Science › Ecology › Wildlife Ecology and Conservation
  • Life Sciences › Agricultural and Biological Sciences › Animal Science and Zoology › Effects of Environmental Stressors on Livestock
Keywords
Smutsia temminckii, thermoregulation, food shortages, climate change, diurnal activity, heterothermy, semi-arid environments, Kalahari, biologging, animal welfare, threatened mammals

About these checks

Registry, retraction, Cites and statistics results are computed, not judged, with one exception: a printed reference that no index lists is read once by a model before it counts as a citation error. Claims are extracted by a model and kept only when their quote appears verbatim in the text; each cited source is then checked, with a model’s help, for whether it says what the claim relies on. Those verdicts can be wrong — read the claim’s quote and the cited source before relying on one.

Claims read by deepseek-v4.1-flash under definition version 3 (extract-v3) on 2026-10-29.

Classified by jev-1.13-20260917 (gold-guideline-v0.2) on 2026-10-29.

Cited sources are checked by gpt-6-sol.

Statistics and arithmetic are recomputed, not judged.

Confidence bands are shown only for fields calibrated on our annotated test set. None is calibrated yet, so no band is shown.

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