Present and past changes in continental weathering and ocean circulation from radiogenic Nd, Hf and Pb isotopes

The past and present variability of ocean circulation and weathering inputs can be reconstructed from the geochemical composition of seawater and marine sediments. Among other proxies, the radiogenic isotope compositions of neodymium (Nd), hafnium (Hf) and lead (Pb) have been applied to reliably trace variations in the ocean that have been related to climate change. However, not all mechanisms and processes governing the geochemical cycling and redistribution of these isotopes are understood in sufficient detail. In order to improve the understanding of some of these mechanisms, the subject of the studies presented in this thesis are the evolution of the radiogenic isotopes of Nd, Hf and Pb in the hydrosphere from the first release into solution during continental weathering via the cycling in present day seawater to the marine sediments, where the information about variations in past ocean circulation and weathering inputs is stored

Glacial reduction of AMOC strength and long term transition in weathering inputs into the Southern Ocean since the Mid Miocene: Evidence from radiogenic Nd and Hf isotopes

Combined seawater radiogenic hafnium (Hf) and neodymium (Nd) isotope compositions were extracted from bulk sediment leachates and foraminifera of Site 1088, ODP Leg 177, 2082 m water depth on the Agulhas Ridge. The new data provide a continuous reconstruction of long and short-term changes in ocean circulation and continental weathering inputs since the Mid-Miocene. Due to its intermediate water depth the sediments of this core sensitively recorded changes in admixture of North Atlantic Deep Water (NADW) to the Antarctic Circumpolar Current (ACC) as a function of the strength of the Atlantic Meridional Overturning Circulation (AMOC). Nd isotope compositions (εNd) range from -7 to -11 with glacial values generally 1 to 3 units more radiogenic than during the interglacials of the Quaternary. The data reveal episodes of significantly increased AMOC strength during late Miocene and Pliocene warm periods whereas peak radiogenic εNd values mark a strongly diminished AMOC during the major intensification of Northern Hemisphere Glaciation near 2.8 Ma and in the Pleistocene after 1.5 Ma. In contrast, the Hf isotope compositions (εHf) show an essentially continuous evolution from highly radiogenic values of up to +11 during the Miocene to less radiogenic present day values (+2 to +4) during the late Quaternary. The data document a long-term transition in dominant weathering inputs, where inputs from the South America are replaced by those from Southern Africa. Moreover, radiogenic peaks provide evidence for the supply of radiogenic Hf originating from Patagonian rocks to the Atlantic sector of the Southern Ocean via dust inputs

Variable climates lead to varying phenotypes: ‘weird’ mammalian torpor and lessons from non-Holarctic species

Mammalian heterotherms, species that employ short or long periods of torpor, are found in many different climatic regions. Although the underlying physiological mechanisms of heterothermy in species from lower latitudes (i.e. the tropics and southern hemisphere) appear analogous to those of temperate and arctic heterotherms, the ultimate triggers and resulting patterns of energy expenditure and body temperature are often noticeably different. Phenotypic flexibility in the patterns of thermoregulation in non-Holarctic species can be extensive (depending on body condition, environmental parameters and species competition) and the factors responsible for inducing heterothermy are more variable in non-Holarctic species. As well as being a regular adaptation to seasonality, heterothermy can also be employed as a response to unpredictability in environmental parameters and as a response to emergency situations. Non-Holarctic heterotherms also challenge the notion that regular inter-bout arousals during hibernation are obligatory and suggest all that is necessary to maintain proper functioning during hibernation is an occasional passive return to -or maintenance of - a relatively high body temperature. The study of non-Holarctic heterotherms has led to the conclusion that heterothermy must be defined on the basis of mechanistic, physiological parameters, and not solely by body temperature; yet we are still limited in our abilities to record such mechanistic parameters in the field. It is now believed that homeothermy in mammals evolved in hot climates via an ancestral heterothermic state. Similar to extant warm-climate heterotherms, early mammals could have relied mainly on passive body temperature regulation with a capacity for short- to longer-term up-regulation of metabolism when needed. Hibernation, as seen in temperate and arctic species may then be a derived state of this ancestral heterothermy, and the study of torpor in warm climates can provide potential models for the energetics of early mammals

Dormancy

Dormancy or torpor is a widely-recognized behavioral and physiological state of bothanimals and plants that generally indicates inactivity and reduced metabolic rate. It caninvolve very different physiological states in response to a variety of environmentalstimuli, including temperature, water, or food. It can last < 1 day, may occur for a fewconsecutive days, or may last an entire season or even many years. Torpor involvesphysiological changes related especially to body temperature, metabolism and waterbalance. Hibernation is when an organism spends the winter in a state of dormancy; it islong-term multi-day torpor for survival of cold conditions. Estivation is summer dormancy, for survival of hot and dry periods. The general roles of torpor, hibernation or estivation are avoidance of unfavorable or lethal short- or long-term (seasonal) climatic conditions and conservation of energy during this period of inactivity. Seasonaldormancy allows species to exploit ephemeral environments and colonize habitats that would otherwise be unsuitable for growth or survival at certain times of the year. Thereare costs to dormancy and torpor, but the advantages contribute to the fitness of individuals and species that use it

Torpor on Demand: Heterothermy in the Non-Lemur Primate Galago moholi

Hibernation and daily torpor are energy- and water-saving adaptations employed to survive unfavourable periods mostly in temperate and arctic environments, but also in tropical and arid climates. Heterothermy has been found in a number of mammalian orders, but within the primates so far it seems to be restricted to one family of Malagasy lemurs. As currently there is no evidence of heterothermy of a primate outside of Madagascar, the aim of our study was to investigate whether small primates from mainland Africa are indeed always homeothermic despite pronounced seasonal changes in weather and food availability., which inhabits a highly seasonal habitat with a hot wet-season and a cold dry-season with lower food abundance, was investigated to determine whether it is capable of heterothermy. We measured skin temperature of free-ranging individuals throughout the cool dry season using temperature-sensitive collars as well as metabolic rate in captured individuals. Torpor was employed by 15% of 20 animals. Only one of these animals displayed heterothermy in response to natural availability of food and water, whereas the other animals became torpid without access to food and water. are physiologically capable of employing torpor. However they do not use it as a routine behaviour, but only under adverse conditions. This reluctance is presumably a result of conflicting selective pressures for energy savings versus other ecological and evolutionary forces, such as reproduction or territory defence. Our results support the view that heterothermy in primates evolved before the division of African and Malagasy Strepsirhini, with the possible implication that more primate species than previously thought might still have the potential to call upon this possibility, if the situation necessitates it

Ambient Temperature Cycles Affect Daily Torpor and Hibernation Patterns in Malagasy Tenrecs.

Hibernation and daily torpor (heterothermy) allow endotherms to cope with demanding environmental conditions. The depth and duration of torpor bouts vary considerably between tropical and temperate climates, and tropical hibernators manage to cope with a wider spectrum of ambient temperature (Ta) regimes during heterothermy. As cycles in Ta can have profound effects on activity and torpor patterns as well as energy expenditure, we examined how these characteristics are affected by daily fluctuating versus constant Ta in a tropical hibernator, the lesser hedgehog tenrec (Echinops telfairi). Throughout the study, regardless of season, the tenrecs became torpid every day. In summer, E. telfairi used daily fluctuations in Ta to passively rewarm from daily torpor, which led to synchrony in the activity phases and torpor bouts between individuals and generally decreased energy expenditure. In contrast, animals housed at constant Ta showed considerable variation in timing and they had to invest more energy through endogenous heat production. During the hibernation season (winter) E. telfairi hibernated for several months in constant, as well as in fluctuating Ta and, as in summer, under fluctuating Ta arousals were much more uniform and showed less variation in timing compared to constant temperature regimes. The timing of torpor is not only important for its effective use, but synchronization of activity patterns could also be essential for social interactions, and successful foraging bouts. Our results highlight that Ta cycles can be an effective zeitgeber for activity and thermoregulatory rhythms throughout the year and that consideration should be given to the choice of temperature regime when studying heterothermy under laboratory conditions

Disparate roost sites drive intraspecific physiological variation in a Malagasy bat

Many species are widely distributed and individual populations can experience vastly different environmental conditions over seasonal and geographic scales. With such a broad ecological reality, datasets with limited spatial and temporal resolution may not accurately represent a species and could lead to poorly informed management decisions. Because physiological flexibility can help species tolerate environmental variation, we studied the physiological responses of two separate populations of Macronycteris commersoni, a bat widespread across Madagascar, in contrasting seasons. The populations roost under the following dissimilar conditions: either a hot, well-buffered cave or within open foliage, unprotected from the local weather. We found that flexible torpor patterns, used in response to prevailing ambient temperature and relative humidity, were central to keeping energy budgets balanced in both populations. While bats’ metabolic rate during torpor and rest did not differ between roosts, adjusting torpor frequency, duration and timing helped bats maintain body condition. Interestingly, the exposed forest roost induced extensive use of torpor, which exceeded the torpor frequency of overwintering bats that stayed in the cave for months and consequently minimised daytime resting energy expenditure in the forest. Our current understanding of intraspecific physiological variation is limited and physiological traits are often considered to be fixed. The results of our study therefore highlight the need for examining species at broad environmental scales to avoid underestimating a species’ full capacity for withstanding environmental variation, especially in the face of ongoing, disruptive human interference in natural habitats. SUPPLEMENTARY INFORMATION: The online version contains supplementary material available at 10.1007/s00442-021-05088-2

Nonshivering thermogenesis in the African lesser bushbaby, Galago moholi

The capacity for nonshivering thermogenesis (NST) plays an important role during arousal from torpid states. Recent data on heterotherms inhabiting warmer regions, however, suggest that passive rewarming reduces the need of metabolic heat production during arousal significantly, leading to the question: to what extent do subtropical or tropical heterotherms depend on NST? The African lesser bushbaby, Galago moholi, enters torpid states as an emergency response only, but otherwise stays normothermic throughout the cold and dry winter season. In addition, this species shows unusual rewarming difficulties during arousal from torpor on cold days. We therefore examined the seasonal adjustments of the capacity for NST of naturally acclimatized G. moholi by stimulation with noradrenaline (NA) injection. Dissection of two adult female bushbabies revealed that G. moholi possesses brown adipose tissue, and NA treatment (0.5 mg kg−1, s.c.) induced a significant elevation in oxygen consumption compared with control (saline) injection. However, the increase in oxygen consumption following injection of NA was not significantly different between winter and summer. Our results show that the ability to produce heat via NST seems to be available throughout the year and that G. moholi is able to change NST capacity within a very short time frame in response to cold spells. Together with results from studies on other (Afro-)tropical heterotherms, which also indicate low or even absent seasonal difference in NST capacity, this raises the question of whether the definition of NST needs to be refined for (Afro-)tropical mammals

Saving energy via short and shallow torpor bouts

Maintaining a high and stable body temperature as observed in most endothermic mammals and birds is energetically costly and many heterothermic species reduce their metabolic demands during energetic bottlenecks through the use of torpor. With the increasing number of heterotherms revealed in a diversity of habitats, it becomes apparent that triggers and patterns of torpor use are more variable than previously thought. Here, we report the previously overlooked use of, shallow rest-time torpor (body temperature >30 °C) in African lesser bushbabies, Galago moholi. Body core temperature of three adult male bushbabies recorded over five months showed a clear bimodal distribution with an average active modal temperature of 39.2 °C and a resting modal body temperature of 36.7 °C. Shallow torpor was observed in two out of three males (n = 29 torpor bouts) between June and August (austral winter), with body temperatures dropping to an overall minimum of 30.7 °C and calculated energy savings of up to 10%. We suggest that shallow torpor may be an ecologically important, yet mostly overlooked energy-saving strategy employed by heterothermic mammals. Our data emphasise that torpor threshold temperatures need to be used with care if we aim to fully understand the level of physiological plasticity displayed by heterothermic species

Limited Physiological Compensation in Response to an Acute Microclimate Change in a Malagasy Bat

Rapid environmental changes are challenging for endothermic species because they have direct and immediate impacts on their physiology by affecting microclimate and fundamental resource availability. Physiological flexibility can compensate for certain ecological perturbations, but our basic understanding of how species function in a given habitat and the extent of their adaptive scope is limited. Here we studied the effect of acute, experimental microclimate change on the thermal physiology of two populations of the widespread Malagasy bat, Macronycteris commersoni. Populations of this species are found roosting under contrasting conditions, i.e., in a constant hot and humid cave or below foliage unprotected from fluctuations in ambient conditions. We exposed free-ranging individuals of each population to the respective opposite condition and thus to novel microclimate within an ecologically realistic scope while measuring metabolic rate and skin temperature. Cave bats in forest setting had a limited capacity to maintain euthermia to the point that two individuals became hypothermic when ambient temperature dropped below their commonly experienced cave temperature. Forest bats on the other hand, had difficulties to dissipate heat in the humid cave set-up. The response to heat, however, was surprisingly uniform and all bats entered torpor combined with hyperthermia at temperatures exceeding their thermoneutral zone. Thus, while we observed potential for flexible compensation of heat through “hot” torpor, both populations showed patterns suggestive of limited potential to cope with acute microclimate changes deviating from their typically occupied roosts. Our study emphasizes that intraspecific variation among populations could be misleading when assessing species’ adaptive scopes, as variation may arise from genetic adaptation, developmental plasticity or phenotypic flexibility, all of which allow for compensatory responses at differing time scales. Disentangling these mechanisms and identifying the basis of variation is vital to make accurate predictions of species’ chances for persisting in ever rapidly changing habitats and climates