Studia Chiropterologica

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1 Studia Chiropterologica Annals of the Chiropterological Information Center Multifactor Analysis of Refugioclimate in Places of Hibernation of Chosen Bat Species Grzegorz Kłys volume 8 (2013) Publications of the Chiropterological Information Center Institute of Systematics and Evolution of Animals Polish Academy of Sciencies in Kraków

2 Chiropterological Information Center Institute of Systematics and Evolution of Animals, Polish Academy of Sciences in Kraków Sławkowska 17, Kraków, Poland Phone: (48) (prefix) (12) , ; Fax: (48) (prefix) (12) , Babiogórski Park Narodowy Zawoja 1403, Poland Phone: (48) (prefix) (33) Fax: (48) (prefix) (33) Printed in Poland by: Zakład Poligraficzno Wydawniczy PLIK Bytom ul. Siemianowicka 98 ISBN Editor-in-Chief Editorial board Scientific Advisory Board Bronisław W. Wołoszyn Tomasz Pasierbek Katarzyna Kozakiewicz Prof. Wiesław Bogdanowicz, Ph.D., D.Sc. (Poland) Prof. Zbigniew Głowaciński, Ph.D., D.Sc. (Poland) Prof. Dumitru Murariu, Ph.D., D.Sc. (Romania) Józef Omylak, M.Sc., Ing. (Poland) Krzysztof Piksa. Ph.D. (Poland) Zoltan Nagý, Ph.D. (Romania) Jordi Serra-Coba, Ph.D. (Spain) Andriy Taras-Baszta, Ph.D. (Ukraine) Reviewers: Prof. Krzysztof Cena Ph.D., D.Sc. (Australia-Poland); Prof. Bronisław W. Wołoszyn Ph.D., D.Sc. (Poland), This issue was edited with financial support from the Babiogórski National Park. Materials published in Studia Chiropterologica may be reproducted only when the source publication is given. Example of literature citation: Kłys G., Hebda M Efect of type of wood used to construct bat boxes. Studia Chiropterologica, 6:

3 Table of contents: Abstract 1. Introduction Purpose of the work Current state of research Ecology and biology of wintering of bats The phenomenon of hibernation and its meaning The phenomenon of hibernation in bats Choice of place of hibernation Migration or hibernation? Characteristic features of refuges The notion of ecoclimate microclimate of an underground system and refugioclimate Characteristics of physical elements which have influence on hibernation of bats Material and methods General Remarks Places of research General overview of devices and methodology of measurement of hibernation environment (refugioclimate) Air temperature (T) Substrate temperature Air flow velocity (v) Relative humidity of air (Rh) Thermal conduction of materials (λ) Air pressure (p) Level of cooling (K a ) Wintering strategies Thermal comfort of a hibernating bat Conduction Radiation Convection Vaporization

4 8. Choice of place of hibernation Introduction Whether there is a relation of choosing by bats higher temperature at first rather than in later months of hibernation Joint or separate analysis of physical factors of refugioclimate Comparison of physical factors (T s ; Rh; v; λ) of refugioclimate of chosen bat species Analysis of particular species Lesser Horseshoe Bat Rhinolophus hipposideros Western Barbastelle Barbastella barbastellus Daubenton s Bat Myotis daubentonii Natterer s Bat Myotis nattereri Greater Mouse-eared Bat Myotis myotis Brown Long-eared Bat Plecotus auritus Summary Air temperature Relative humidity of air Air flow velocity Thermal conduction Air pressure Wintering strategies Standard of conditions of measurement Synthesis of chosen physical factors Theoretical and practical meaning What is new Literature cited Appendix List of acronyms and symbols used in the work Glossary

5 Abstract There is an enormous number of publications concerning ecology of wintering of bats, but complex research on the choice of place of wintering by bats was not performed so far. Few works from this scope deal with analyses of single parameters, rarely they concern larger number of parameters. It is generally known that rules of transferring heat are governed by laws of physics. There are four ways of transferring heat: conduction, convection, radiation and vaporization. The author in his research attempts to integrate particular abiotic factors in the immediate place of wintering of bats which is called refugioclimate. The following quantities were taken into consideration in the work: air flow velocity (v), air pressure (p), relative humidity (Rh), thermal conductivity of the substrate on which bats wintered (λ) and air temperature (T a ). Six most common in underground systems bat species were selected for the research: lesser horseshoe bat Rhinolophus hipposideros, greater mouse-eared bat Myotis myotis, Daubenton's bat Myotis daubentonii, Natterer's bat Myotis nattereri, brown long-eared bat Plecotus auritus and Western Barbastelle Barbastella barbastellus. Due to the fact that values of physical quantities are often relatively diversified in underground systems a methodology of measurement of physical factors which influence wintering of bats was established. Measurements were performed in the immediate proximity of a wintering bat. Six basic strategies of wintering of bats were proposed depending on their share in cooling a bat. The author of the current work made an attempt of a comprehensive evaluation of the influence of sum of chosen factors on the choice of hibernation place. Additionally, two basic questions were posed: are the chosen abiotic factors of refugioclimate: T s, Rh, v, λ responsible for the choice of hibernation places? Do factors of refugioclimate differentiate species and strategies as far as places of their choice is concerned? It was assumed, that statistical analyses should facilitate evaluation and interpretation of obtained results. Therefore an attempt to answer the following questions was made: Do bats choose higher air temperature at the beginning of hibernation? Is there a relation in the choice of physical factors T s ; Rh; v? Should each of the aforementioned factors be examined jointly or separately? Are there any differences in choice of place of hibernation by 6 chosen bat species in relation to abiotic factors of physical parameters of refugioclimate (T s ; Rh; v)? In order to evaluate influence of the sum of these factors on the choice of hibernation place the author performed measurements of temperature (T a - n=6389), relative humidity (Rh - n=6389), air flow (v - n=6389) and atmospheric air pressure (p - n=6389) of refugioclimate. Only part of results of physical values of refugioclimate of chosen six species of bats was used for the analysis. It produced in total items of data. 5

6 It has been proved that the choice of hibernation place of a given bat species and its strategy is dependant on physical conditions of refugioclimate such as T s ; Rh; v and thermal conductivity of substrate (λ) on which a bat hibernates. It has been demonstrated that each of the examined species possesses its own scope of physical quantities of a refuge which is optimal during hibernation, which quantities supplement one another and which differ one species from another. Wet Kata cooling power degrees were used in order to obtain single values of the analysed parameters such as T s ; Rh; v. For the first time empirical equations of three variables have been made which determine choice of hibernation places and strategies for analysed species. The obtained results and conclusions from the current paper allow to present two important theoretical and practical meanings in ecology and protection of bats in various ecosystems. They are: Comparison of factors which condition choice of a hibernation place may have significant meaning in practice of protection of bats and their habitats. There is a possibility of simulation of microclimate of an underground system and consequently a refugioclimate to the needs of hibernation of a given bat species. 6

7 Introduction Bats are second after rodents group of mammals which is the most common around the world. Thanks to their ability of active flight they are able to carry out longdistance travels during seasonal migrations (Calisher et al., 2006). Flight ability of these animals requires intensive metabolism, high body temperature and significant level of oxygen consumption (Altringham, 1996). Thanks to the phenomenon of heterothermy bats are able to manage sparingly energy from reserves gathered during the period of activeness. Heterothermy allows to last long seasonal drop in external temperature and lack of food in the state of hibernation. In this period at values of body temperature close to ambient temperature metabolism of bats is reduced to minimal values (Hock, 1951; Dunbar and Tomasi, 2006; Boyles et al., 2008). The ability to hibernate is one of key aspects of settling high latitudes (both northern and southern) by bats. In unfavourable season bats in this climatic zone utilize possibility to shelter in various kinds of underground systems. They are used mainly for hibernation, because in winter period they provide sufficient set of physical conditions essential to survive this period without food. Research on bats is complicated due to methodological difficulties connected with their specific way of life, so collected information is often a result of accidental observations. The most intensive research on this group took place in the second half of the 20th century in Europe, North America and some tropical countries (Kunz et al., 1983; Altringham, 1996; Neuweiler, 2000; Evelyn, Stiles, 2003; Kunz, ter Hofstede Fenton, 2003; ter Hofstede, Fenton, 2005; Chaverri et al., 2007; Chaverri et al., 2007). Despite numerous research the knowledge of bats still includes lots of blank pages. So far, research focused mainly on measurements of temperature and humidity, very rarely on air flow (Kłys, 2004). Getting to know the most important factors of habitat which determine starting winter sleep (hibernation) by bats and getting to know relations between determinants of habitat and ecoclimate may have significant meaning both theoretical and practical one. Knowledge of these relations may allow to create artificial habitats for species of bats which are endangered and dying out; it also creates possibility to model conditions of refugioclimate for the needs of hibernation. Currently, attempts are made to protect and manage underground systems which take into account needs of bats (Mitchell-Jones et al., 2007). However, so far many of the conditions which determine settlement in underground systems and distribution of bats therein have not been defined. 7

8 Acknowledgments The author wishes to thank Prof. Jerzy Lis, Ph.D., D.Sc. for his support and help during work. I am grateful to the staff of the Faculty of Mining and Geology, Institute of Mining, at the Silesian University of Technology in Gliwice for scientific help: Paweł Wrona, Ph.D., Ing., Zenon Różański, Ph.D., Ing., Grzegorz Pach, Ph.D., Ing. I express my gratitude to Andrzej Sobolewski Ph.D., Ing. from Central Institute for Labour Protection National Research Institute in Warsaw; Jacek Piasecki Ph.D. from the Department of Climatology and Atmosphere Protection, Department of Earth Sciences and Environment Shaping the University of Wroclaw; Grzegorz Wojtaszyn Ph.D. of the Polish Society for Nature Protection Salamandra ; Magdalena Dziegielewska Ph.D. of Applied Entomology Department of the Agricultural University in Szczecin; Marek Bandrowski of the Association of Friends of Police "Treasure ; the staff of Moravský Kras Landscape Protection Park (Chráněná krajinná oblast ChKO Moravský Kras) in the Czech Republic, in particular to Mr Miroslav Kovařik for enabling research of lesser horseshoe bat Rhinolophus hipposideros; the Ministry of Environment; Provincial Nature Conservation Authorities in Opole, Katowice, Wrocław, Gorzów Wielkopolski, Szczecin and Lublin. I also thank Mr Paweł Rutkowski from FLIR Systems Inc. for taking infrared pictures of wintering bats, as well as all other people who made this work possible. Thanks are also to Grzegorz Gurbała, who translated this paper into English. 8

9 1.1. Purpose of the work The main purpose of this work is an attempt to determine and evaluate influence of abiotic factors on the choice of hibernation places by chosen species of bats in cave environment and in anthropogenic facilities taking as an example various underground systems. Earlier research did not take into consideration if the choice of hibernation place as well as the strategy of this choice are dependent on physical parameters of refugioclimate, and simultaneous influence of its three basic parameters (T, Rh, v) (see: List of acronyms and symbols used in the work) was not analysed, as well as influence of the substrate on which bats winter (λ degree of thermal conductivity). The author of the current work made an attempt of a comprehensive evaluation of the influence of sum of these factors on the choice of hibernation place. Additionally, two basic questions were put: Are chosen abiotic factors of the refugioclimate: Ts, Rh, v, λ responsible for the choice of hibernation places? Do factors of refugioclimate differentiate species and strategies as far as places of their choice is concerned? It was assumed, that statistical analyses should facilitate evaluation and interpretation of obtained results. Therefore an attempt to answer the following questions was made: Do bats choose higher air temperature at the beginning of hibernation? Is there a relation in the choice of physical factors Ts; Rh; v? Should each of the aforementioned factors be examined jointly or separately? Are there any differences in choice of place of hibernation by 6 chosen bat species in relation to abiotic factors of physical parameters of refugioclimate (Ts; Rh; v)? 9

10 1.2. Current state of research Only four out of eighteen currently known families of bats, namely Rhinolophidae, Vespertilionidae, Miniopteridae and Molossidae, possess ability to hibernate, which allowed them to settle in the temperate zone. Bats which inhabit temperate and northern climates, where seasonal changes of climatic conditions and lack of food appear, need specific morphological, physiological and behavioural adaptations. For a long time this problem have been arousing particular interest among scientists (Panugajewa, Slonim, 1953; Punt, van Nieuwenhoven, 1957; Kalabukhov, 1985; Tiunow, 1988; Ransome, 1990; Arnold, 1993; Koteja et al., 2001; Humphries et al., 2002; Dawydow, 2004; Geiser, 2004; Anufrijew, 2005; Anufrijew, Rewin, 2006). The ability to survive in low temperatures of various animal species is known since the 19 th century. Professor A. Horvath (1878) observed drop in body temperature of striped ground squirrels to 0,2 C. In 1912 benefit from decreasing body temperature was demonstrated experimentally by P.I. Bahmjetjeba (in: Kalabukhow, 1985). It was stated then that in the state of hypothermia body temperature may drop to -9 C without creating ice in body fluids. Research conducted later showed that minimal temperatures to which an animal may be cooled without the danger of creating ice in tissues are different in various species and in general they are lower at slowed cooling (Kalabukhow, 1985). During recent years on the basis of experimental and field research basic knowledge of the phenomenon of hibernation in various groups of vertebrates and invertebrates was acquired (Kalabukhow, 1985; Pastuhow, 1986; Slonim, 1986; Hoczaczka, Somiero, 1988; Geiser, Ruf, 1995; Geiser, 2001, 2004а, b; Janicki, Cygan- Szczegielniak, 2006; Cooper, Geiser, 2008; Wojciechowski et al., 2011). These animals have a common feature: their organisms stop using up external supply of food after they enter the state of hibernation and at the same time in various, but always a controlled way, they reduce intensity of metabolism (Hoczaczka, Somiero, 1988). Hibernation allows to reduce the use of energy by 95% in comparison to the costs of keeping activeness during winter and preserving energy is correlated with temperature of hibernation (Geiser, 2004а). The process is not homogeneous, it is broken periodically by awakenings of the animals which constitutes a significant part of costs of energy of an organism in winter period (up to 84%, Boyles et al., 2008). For some species of bats calculations of the budget of time and energy in the state of hibernation were performed, in particular for a North American species of little brown 10

11 bat Myotis lucifugus (Thomas et al., 1990) and an Eurasian species of Daubenton s bat Myotis daubentonii (Matveev et al., 2005). In Myotis lucifugus 15 days-long cycles of awakenings were observed. So during one cycle it uses up about 3.6 kj which corresponds to energy contained in approximately 0.1 g of fat (Thomas, 1995). In this species there are on average from 10 to 13 such cycles. However, other research show that duration time of one cycle of lethargy readiness of bats may differ significantly from several days to several months (Strielkow, 1971b; Geiser, 2004а). Duration time of a cycle may depend on external factors (temperature in a winter refuge, humidity, change in atmospheric pressure), species, sex and individual features of a specimen (Strielkow, 1971b; Park et al., 2000). Since energy losses during awakenings constitute a significant part of wintering energy budget (Thomas et al., 1990), causes and functions of such unnecessary behaviour of bats aroused and still arouse interest. Hypotheses which try to explain this phenomenon (Park et al., 2000) may be divided into two groups. I. Awakenings caused by abiotic factors: Searching for optimal temperature for hibernation (Boyles et al., 2006). Compensation of lost moisture while breathing and evaporation through skin in refuges of insufficient humidity (Thomas, Cloutier, 1992; Thomas, Geiser, 1997). II. Awakenings caused by biotic factors: Getting rid of by-products of metabolism (Park et al., 2000). Reproduction (Boyles et al., 2006). Acquiring food (important for areas with mild climate) (Avery, 1985; Brigham, 1987). Abiotic factors in a refuge not only may have on frequency of awakenings of the animals, but also determine risk of survival during hibernation. So far works have taken into consideration only temperature and humidity as factors which have influence on conditions of wintering (Nagel, Nagel, 1991; Visnovska et al., 2006;. Boyles et al., 2008; Boratyński et al., 2012). Optimal temperature for most bat species which winter in underground systems of the temperate zone equals from 0 to +10 C (Webb et al., 1995; Humphries et al., 2002). Awakenings are connected with searching for optimal temperature in winter refuge in order to maximize saving energy (Speakman, Racey, 1989). 11

12 Research on energy expenses during hibernation of mammals often emphasise physiological aspects of thermoregulation, but do not take into account behaviour of mammals (Humphries et al., 2002). For instance, to form dense groups (aggregations) animals use euthermia in order to reduce heat loss, but it may also influence use of energy during hibernation of mammals. Aggregation is a phenomenon belonging to important physiological and ecological behaviours for hibernating animals (Boyles et al., 2008). Creating groups during hibernation is changeable within a species and between species. A tendency is observed to aggregate even in the same winter refuge. Some bat species form small, thin clusters, whereas other species create dense groups (Smirnow et al., 1999; Tomilienko, 2002) which contain up to a few thousands of specimens (Betke et al., 2008). During winter bats can move and change location, which leads sometimes to change in size and composition of aggregation in a hibernaculum (Orlowa et al., 1983; Tomilienko, 2002). It is believed that it is possible to form dense concentrations which allow to compensate loss of metabolic heat to the environment (Boyles et al., 2008). Body temperature of animals in the state of hibernation usually fluctuates around ambient temperature (Geiser, 2004a). However, research on the phenomenon of hibernation is usually limited to descriptive field observations (e.g. when and where mammals wintered) or physiological research in a laboratory (e.g. testing metabolism and internal secretion, functions of animals) which have no environmental context. Humidity of a wintering place has large influence on duration time of the lethargy readiness cycle. Influence of this parameter is often more important than nutritional status of an animal (Thomas, Cloutier, 1992; Thomas, 1995). Even in conditions of high relative humidity (90-98%) the loss of moisture may be significant. The loss of moisture occurs through external surface of the animal s body and through breathing. Loss of water from an organism that is too high stops normal course of metabolic processes and may lead to death. This is why in refuges of lower humidity bats often wake in order to replenish water in their organisms. Resilience to loss of water in various species is different (Ransome, 1990). Research conducted on Pipistrellus pipistrellus shows that animals wake up to replenish water in their organisms caused by vaporization (Speakman, Racey, 1989). By creating dense aggregations animals will reduce water loss and probably they will wake up less often, which will lead to a reduction of energy expenses in winter period (Ransome, 1990; Thomas, Cloutier, 1992; Thomas, 1995). Research conducted so far state that humidity and temperature may have influence on choice, duration time of the cycle of awakenings and formation of clusters of bats. It is obvious that an organism of a hibernating bat is influenced by a whole set of 12

13 factors (not always known). This is why data collected from hibernation places should concern above else cooling factors such as air flow velocity, temperature and thermal conductivity of the rock, air temperature, relative humidity and wintering strategy. The author stated that air flow velocity is one of the most important factors which condition bat hibernation (Kłys et al., 2002; Kłys, 2004, 2008). One of the issues in research on hibernation should be estimation of value of particular physical factors which complement each other. Equally important is methodology of measurement of these factors (Kłys et al., 2005). Many works concerning meteorological conditions in a place of occurrence of bats lack the description of methodology of measurement and devices used, which makes interpretation of such data impossible (Kłys et al., 2005). Influence of presence of the person performing measurements on their accuracy was also neglected (Kłys et al., 2005). Despite the fact of existence of an enormous number of publications concerning ecology of wintering of bats, complex research on the choice of place of wintering by bats was not performed. Few works from this scope deal with analyses of single parameters. Kłys, Wołoszyn, (2005) proposed a term of refugioclimate for description of the set of physical factors which have an effect directly in hibernation place of a bat. The author in his research attempts to connect particular abiotic factors in an immediate place of wintering called a refugioclimate. 13

14 14

15 2. Ecology and biology of wintering of bats 2.1. The phenomenon of hibernation and its meaning In order to achieve high level of independence from restrictions imposed on them by environment, the endothermic organisms developed ability to create internal warmth to keep high and stable body temperature (Smith-Nilsen, 2008). However, this ability requires high energy costs. In many animals, especially small ones, costs connected with thermoregulation may exceed the amount of available energy. In order to solve the problem of shortage of energy, they enter a state of torpor. This strategy allows them to avoid energy-intensive, fast pace of metabolism necessary to maintain stable body temperature. It is a precisely regulated and controlled physiological state. Torpor is then an answer to unfavourable environmental conditions (shortage of food, water, cold, warmth) and is characterised with drastic drop in body temperature and other physiological functions. This state may last from several hours to several months Hoffman, 1964; Lyman (Hoffman, 1964; Lyman et al., 1982; Nelson, 1980; Wang, 1987; French, 1988; Storey and Storey, 1990; Geiser and Ruf, 1995). In contrast to ectothermic organisms (e.g. amphibians and reptiles), endothermic organisms are able to leave the state of torpor at any moment. With the use of thermogenesis they bring back normal body temperature. Representatives of many groups of mammals have an ability to enter the state of torpor. In the majority they are small animals, characterised with high metabolism, for which maintaining stable body temperature with shortage of food at the same time is to costly in terms of energy, for instance Tachyglossidae from Monotremata many Dasyuridae from Marsupialia, Xenarthra, Afrosoricida, Crocidurinae and Erinaceidae from Insectivora, many bat species of both suborders Megachiroptera and Microchiroptera, small Primates and various Rodentia: Cricetidae, Heteromyidae, Muridae, and Sciuridae, Carnivora: (Ursidae), and some Mustelidae, from Canidae Nyctereutes. Apart from numerous species of mammals there are only a few bird species known such as Phalaenoptilus nuttallii several species from the family of Trochilidae, Apodidae and mousebirds from Colius genus which enter the state of torpor (Lyman, 1982; Geiser, 2001; 2004a; Schmidt-Nielsen, 2008). Torpor in homoiothermal organisms may be characterized depending on duration time, depth of that state and as seasonal or non-seasonal. Seasonal torpor may be accomplished through estivation and hibernation; it is characterized with states of torpor which may last several hours, days, weeks or months within the period of one season. 15

16 There are two forms of torpor: a short one, called daily torpor, and a long one, called extended torpor or hibernation. Organisms which are able to enter into the state of hibernation are called hibernators. In reality seasonal torpor never encompasses entire season of hibernation, but it is broken by temporary awakenings and short periods of normothermia (French, 1985, 1988). There are many different kinds of torpor which differ in frequency of heartbeat, breath and body temperature. It results from a fact that torpor formed independently in various phyletic lineages of mammals and is caused by various ecological determinants, which in turn lead to enormous variety of patterns of torpor (Geiser, 1988). Due to that some forms of torpor are called differently and borders between these forms are fuzzy. In relation to animals which enter into the state of torpor in winter, the following terms are often used: winter sleep, winter rest, winter lethargy or hibernation. In some species during the rest thermoregulation is not stopped, body temperature drops minimally and the animals do not leave their winter refuges and do not feed. In each of these cases the purpose is to achieve hypometabolic state to save energy. The state of torpor usually consists of cycles: entering, maintaining and awakening from torpor. Many hormonal changes occur in a body of the animal. Entering the state of torpor is characterized by the following features: significant drop in metabolism, slowing down heartbeat down to several beats a minute, narrowing blood vessels, reduction of pace of breathing, including occurrence of apnea, drop in oxygen consumption, significant drop in body temperature, often to the level of slightly higher than ambient temperature, decrease in brain activity, decrease in nervous excitability which manifests itself in body torpor and faint reaction to external stimuli. All physiological functions are reduced to a minimum. Despite large-scale research on winter sleep of animals, basic natural mechanisms of hypometabolism are still hardly known. 16

17 2.2. The phenomenon of hibernation in bats In the temperate climatic zone the sustenance of bats are insects and other arthropods. Insects show significant fluctuations of numerical strength throughout the year. In cool season of the year when there is lack of food bats must migrate or limit drastically the use of energy to survive (Fig. 1). To a great degree the specificity of thermoregulation and energetics of bats is determined by the type of food and geographical distribution. Taking into account ability to regulate body temperature (Tb) depending on the influence of ambient temperature (Ta) bats may be artificially divided into three groups (Davydov, 2004): definite homoitherms relative homoitherms, which are able to enter into torpor homoitherms which are able to regularly enter into torpor and are able to hibernate. Specific feature of hibernation is switching off thermoregulation. The external stimulus which provokes the process of change is above else lowering ambient temperature. Basic strategy of hibernating bats is searching for an appropriate refuge (hibernaculum) in which ambient temperature does not drop below 0 C, appropriate humidity and air flow velocity is preserved and the animal is hidden from predators. Threshold temperature is different for particular species. Above the threshold temperature hibernation can not occur. In hibernation typical for bats body temperature drops within minutes, depending on the size of a bat and its fat reserves, to about several C for the period of from a few to a dozen or so weeks, which is the precondition to survive without feeding. Body temperature is then kept around the ambient temperature (Fig 2). Energy to live is obtained from fat reserves accumulated for this purpose. Bats produced winter sleep glands which are characteristic for them, i.e. brown adipose tissue which accumulates fat of high energetic value. Energy contained in it is used for fast heating up the body during awakening, when surrounding conditions exceed threshold value. If ambient temperature rises, an animal returns to regular activity, whereas drop in temperature below 0 C threatens the organism with freezing, which is why the animal increases pace of metabolism to maintain its body temperature on the level slightly above 0 C or awakes to replenish fat reserve or change refuge. Awakening may occur also under influence of other strong external stimuli. Large role during hibernation is played by social thermoregulation. Since most of species hibernate in aggregations, while they sleep they cuddle up to one another which reduces heat loss (Ransome, 1990). Larger bats have lower ratio of space to volume than small ones. Therefore they will use up their energy reserves slower than smaller ones. 17

18 Fig. 1. Annual cycle of activeness of bats from the temperate zone. Fig. 2. Wintering greater mouse-eared bats Myotis myotis. Infrared photographs (FLIRSYSTEM for the author). 18

19 2. 3. Choice of place of hibernation Migration or hibernation? For many organisms which live in the zone of temperate climate, including bats, winter is a critical period. Only proper adaptation to low temperatures in the period of lack of sustenance allows bats to survive in such conditions. Among strategies of survival one may enumerate: transition to an alternative and more substantial type of food, storing food in the times of abundance, migration to a place where food can be found entering the state of hibernation. Most of bats choose the last strategy, which may be called an escape in time in contrast to the strategy of: an escape in space concerning migration of many species of birds (Wołoszyn, 2007, 2008). Like in many other animals also in bats there are sedentary, nomadic and migrating species. In Central Europe species are considered sedentary if they wander no further than km. This group of bats includes among others genera Rhinolophus, Plecotus and also some small species of the genus Myotis. There are also such species which regularly change place of stay from 100 to several hundred km. They require appropriate underground systems for wintering. They are among others: pond bats Myotis dasycneme and greater mouse-eared bats Myotis myotis. Bats from the group of migrating ones cover annually a distance of over a thousand kilometres. Species of the genus Nyctalus, the species of Nathusius pipistrelle Pipistrellus nathusii and parti-coloured bat Vespertilio murinus belong here. In some migrating species it happens that local population has a convenient hibernaculum at their disposal; in that case they resign from migration. When bats migrate they not always choose south direction, which means a warmer zone, but sometimes also north to a known hibernaculum. It is often seen as a precondition of colonization of areas of low temperatures (Raphae et al., 2000). Hence hibernation seems to be the best solution for bats of the temperate climatic zone Characteristic features of refuges Availability of observation of hibernating bats is varied. Most of data comes from caves and underground systems in which access of an observer is the easiest one. To the remaining refuges access is limited and data on bats which hibernate there are rather accidental. 19

20 Bats are usually very flexible animals as far as the choice of refuge is concerned. They settle refuges of both natural and anthropogenic origin (Table 1). Table 1. The most frequently found refuges of bats in Poland. Type of refuge Chosen works Natural shelters Caves Kowalski K., 1954; Wołoszyn, In heaps of stones, under the stones in soil, animal burrows Tree hollows, crevices in tree trunks and under the bark Strielkow, 1970; own observations Krzanowski, 1956; Ruprecht, 1976; Postawa et al., Crevices in rocks Vlaschenko, Naglov, 2005; Strielkow, Ilin, 1990; Borissenko et al., Anthropogenic shelters Bunkers, mines, bomb shelters, drain pipes, cellars, sewers and other underground systems Kepel, 2005; Kłys, 1994; Lesiński et al., Wells Kowalski M., 1995; Ignaczak, Radzicki, Cracks in buildings, under window sills Parts of buildings above the ground, ventilation ducts, attics, empty space in walls Lesiński, 2006; Kłys, 1996; Wołoszyn, Kłys, 1996; Krzanowski, 1980; Olszewski, 2003; Iwaniuk, Szkudlarek, Nesting boxes for birds and bats Kłys, Hebda, Timbering, window shutters, gaps between beams own observations Concrete structures, bridges Wojtaszyn et al., 2005; Ciechanowski,

21 Natural shelters Caves and crevices in rocks are used by bats mainly during winter (places of hibernation) and autumn (mating shelters), more rarely during summer. Breeding colonies of bats in Polish caves are created sporadically, however it concerns only two species: greater mouse-eared bat (currently only one colony is known, in Studnisko Cave in Krakowsko-Częstochowska Upland) (Wołoszyn, 2008) and lesser horseshoe bat. These places may also play the role of daytime hiding places for males in spring and summer period. Hiding places in tree hollows and under the bark may be settled by bats in all seasons of the year. Some species prefer cracks in tree trunks and boughs (e.g. barbastelle bat and lesser noctule), while others (e.g. common noctule) prefer tree hollows made by woodpeckers. Noctule bats and pipistrelle bats can winter in hollows of thicker trees. In mating season, males of some species (noctule bats, pipistrelle bats) occupy separate hollows treating them as their mating shelters. Sometimes, they are also shelters of breeding colonies. Practically nothing is known about bats which winter deep in narrow crevices in rocks or which bury themselves below ground level. Artificial shelters (anthropogenic) Anthropogenic shelters are currently the most important refuges for bats in Western Palearctic. Drifts, tunnels, city sewers or wells create similar conditions as caves. They replace caves for bats in areas where no caves are present. Cracks and gaps in walls of buildings and bridges resemble crevices in rocks, similarly gaps in roofs and shutters substitute hiding places in old trees. Anthropogenic shelters are used mainly during winter, as hibernation places, and autumn as mating shelters. Objects of this type are used for wintering by most bat species (Rhinolophus, Myotis, northern bat, Plecotus, Barbastella). Small, home cellars are used for wintering most often by Plecotus and Daubenton s bats (Lesiński, 2006). Wells, which have microclimate like vertical caves, are an important place of hibernation in areas devoid of other, larger underground sites. City sewers as winter habitat of bats were discovered only a dozen or so years ago (Wojtaszyn et al., 2005). Parts of buildings which are above ground level are for some species (particoloured and serotine bats, noctule bats, pipistrelle bats) main winter shelters. Breeding colonies of bats are usually placed in attics of buildings, as well as cracks and gaps in walls and under boarding, or even space between gutters and walls of a building. Some species (e.g. greater mouse-eared bats) require large and spacious 21

22 attics, while others use also smaller shelters and often hide in gaps (between layers of a roof, between walls, etc.). In mating season males of some species may use nooks in attics or crannies in walls of buildings as their mating shelters. All species of bats in Poland were observed among other places in buildings. Nesting boxes for bats and birds are hung in places with few natural shelters. Bats (some species) settle in them mainly during spring, summer and autumn, but rarely in winter, due to poor insulation against external conditions. They may be shelters for entire breeding colonies. Old buildings, especially wooden, like sheds, damp cellars, attics, outbuildings and church towers used to be inhabited by bats on a large scale. Currently, buildings are usually insulated and impenetrable, so bats return to caves and hollows, under bridges and into old wells, any place where microclimatic conditions of the interior are favourable enough for them to hibernate. State of knowledge of wintering places of bats is still unsatisfactory. 22

23 3. The notion of ecoclimate microclimate of an underground system and refugioclimate In research on hibernation of bats (biospeleological, speleoclimatic) there is certain divergence in terminology and interpretation of issues concerning climate of underground systems (caves) between botanists, zoologists and climatologists. It results mainly from divergence in the research subject of those three branches of science. While carrying out research on climate of caves as habitat of animals and plants one should use a term of habitat climate or ecoclimate. This term is presented by B. W. Wołoszyn (1976) and describes a set of meteorological factors which influences given environment, which in this case are underground systems. On the other hand if the purpose of research is to determine a set of climatic factors which occur in underground systems for a period of many years (a year and longer) a term of speleoclimate seems to be more adequate (Skalski, 1973). Microclimatology has been sectioned off as one of the branches of climatology (Whittow, 1986). Just like climate or topoclimate (local climate) it has some distinctive features in relation to surrounding areas (Yoshino, 1975). Wiglej and Brown (1976) present different approach to the subject. According to these authors many speleological issues, among other things connected with climate which occurs inside underground systems, may be reduced to physical problems, which is why they use the term of cave meteorology. In this context the term meteorology is used in a broader sense and includes also climatology. To be precise, this term refers also to dynamic elements of quickly changing atmospheric conditions, while climatology deals with long-term changes. This approach results from a fact that underground systems are dynamic environments and temperature, humidity, air flow, etc. show short and long-term spatial and temporal changeability. In Polish speleological literature the most frequently used term is cave microclimate, as a description of a set of climatic conditions existing in caves, while state of external atmosphere and changes which occur there are examined by topoclimatology and climatology. The term micrometeorology is also used, as it deals with processes of exchange of momentum, heat and matter between atmosphere and its substratum (Paszyński et al., 1999). In meteorological lexicon (Niedźwiedź, 2003) microclimatic conditions of closed spaces (artificial and natural), including caves and animal burrows, are called cryptoclimate (interior microclimate, climate of a closed space). 23

24 In the current work a notion of interior microclimate was adopted as a set of conditions existing inside the examined system. It is considered to be background of values of existing generally particular factors of the underground system. The term refugioclimate used in the work was proposed in 2005 (Kłys, Wołoszyn) for physical conditions in the immediate proximity of hibernation place (several cm around a wintering bat). It results from the fact that measurements of thermal background, air movement and humidity (microclimate of the underground system) were often significantly different from these factors in the immediate proximity of the wintering bat. Many times they are rock niches or recesses while sometimes the examined object hangs centrally in a part of the gallery and then these values do not differ significantly from values of factors of the interior s microclimate. This notion defines the problem correctly and at the same time it refers to the specific values essential for further discussion. It refers to a specific place of hibernation, not to the microclimate of entire or a part of the underground system. Obviously, a microclimate as a set of meteorological factors (local peculiarities of climate) influences directly conditions of a refugioclimate and what follows the existence of the organism. In underground systems, which are often places of hibernation of bats, there are different microclimatic conditions of the interior in relation to the surface conditions. They change already in short distances (Chromow, 1977). Microclimate of an interior is more stabilized and mild. It depends of course on the size of the underground system and the scale of climatic change, long term, seasonal or daily ones. It is usually characterized by lower than ambient temperature and higher humidity, as well as air circulation which changes throughout the year. However in the other, opposite part of the system the changes are reverse to the observed ones. Climatic conditions of an underground system are influenced by height of the entrance opening above sea level, exposure of the opening, existence of flora above the opening, morphology of the cave (length and shape of corridors and chambers), streams of water transferring heat and taking it out, thermal energy from the Earth s interior. 24

25 4. Characteristics of physical elements which have influence on hibernation of bats As heterothermic organisms bats can regulate body temperature within certain limits. Their thermoregulation is so special, that they can lower their body temperature almost to the level of ambient temperature. However, within moments (Davydov, 2004) bats can raise temperature considerably higher than ambient temperature even to 40 C at ambient temperature of just several degrees. So bats of temperate zones enter into torpor in cold season to wait out the period in which there is lack of appropriate sustenance. They lower their body temperature during hibernation almost to ambient temperature (McNab, 1982). It causes significant reduction of metabolism (Thomas, 1995). Independently of research on biological factors of bat hibernation there are also performed microclimatic observations of underground systems. Apart from biological factors, such as ensuring safety from predators, there is a large number of physical factors which have an influence on hibernation. 1. chemical and physical composition of air 2. ability to attach itself to the substrate (Fig. 3) 3. air movement 4. air temperature 5. temperature of surrounding objects (rock) 6. air humidity 7. thermal conduction of rocks 8. phasic changes of air 9. permeating geothermal heat 10. air pressure 11. influence of external conditions In underground systems microclimatic conditions play a very crucial role. They depend on functioning of the entire system. These statements are confirmed by abundant literature devoted to underground microclimate. In case of formation of microclimatic 25

26 conditions of underground systems three closely connected processes bear the essential meaning (Piasecki et al., 2001): exchange of air between the underground environment and its surroundings; flow and exchange of heat between the orogen and air and water in the underground system; circulation of water in the underground system, including circulation of moisture in the form of vaporization and condensation. Through operation of these factors a relation appears between microclimatic conditions in the underground systems and the topoclimate and, in consequence, the refugioclimate (Kłys, 2008). Fig. 3. A roof of an underground system with marked attempts of bats to attach themselves to it. Photographed by the author. Taking into account the set of these factors, one may empirically determine climatic zones for particular species and strategies of bats. In research on hibernation of bats air temperature is often considered to be the most important factor, since it is a universal indicator which reacts quickly for even slight changes in circulation and humidity of underground systems. However, it is very 26

27 hard to examine temperature of a given system without knowledge of direction and intensity of air movement and humidity. Underground environment, including caves, is considered to be stable, which means the one in which both temperature and humidity, as well as other factors are considered to be constant elements. In reality however, these parameters undergo significant changes, though they are not so clear as outside in the open atmosphere. So underground environment has got a dynamic character in which factors show spatial and temporal diversity (Kłys, 2008). One may distinguish many factors which shape temperature inside underground systems. However, the most important ones include: air movement and influence of so called cave winds, which moderate and even out thermal differences between an underground system and its surroundings as well as in the interior of the system. It is accepted that average annual outside temperature corresponds to inside temperature of the underground and decreases as height above sea level grows. While investigating changeability of air temperature profile, the processes of releasing or absorbing latent heat due to condensation and evaporation cannot be ignored. The air that gets inside an underground system usually gets warmer or cooler in contact with a rock. Depending on temperature of the rock being higher or lower than dew point, water vapour condensates or evaporates from the surface of the rock. In both cases during transition of particular amount of substance from one phase into another at constant pressure and temperature, certain amount of heat will be emitted or absorbed, called latent heat or heat of phase transition. However, heat of condensation is equal in terms of absolute value to heat of vaporization, but with opposite sign. Both immediate heat exchange between the rock and the air as well as latent heat absorbed or emitted are very vital in determining air temperature profile in underground systems. Certain amounts of heat (cold) are supplied to underground systems by inflow of external air. They are subjected to quick transformation in underground environment. They take part in determining spatial diversity of course and profile of temperature. In parts which are located deeper amplitude of temperature is small and extreme values of temperature are caused by a more intensive flow of warm or cool air. By absorption and emission of heat rock regulates changes of air temperature caused also by other factors. 27

28 28

29 5. Material and methods 5.1 General Remarks Research was conducted both in caves and artificial shelters. In the first stage an inventory of winter refuges was made, an insight into composition of species was gained and numerical strength of bats was recognized. Next, ecological observations were started, namely examination of structure of groups and interactions between species of wintering bats. This stage of research allowed to accept a working hypothesis that the course of hibernation in bats is significantly and directly influenced by a lot broader spectrum of physical factors than it was believed so far. Six most common in underground systems bat species were selected for the research: lesser horseshoe bat Rhinolophus hipposideros, greater mouse-eared bat Myotis myotis, Daubenton s bat Myotis daubentonii, Natterer s bat Myotis nattereri, brown long-eared bat Plecotus auritus and Western Barbastelle Barbastella barbastellus). Description of places chosen by particular species in winter period concerned temperature, humidity and air flow. Temperature of surface on which bats winter was randomly recorded as well as the type of material on which they winter in order to attribute thermal conductivity. Data was collected mainly in the period of peak of hibernation (December February). Due to the fact that waking bats was avoided, sex and age were not determined, so there is no data if these two factors have influence on the choice of particular place of hibernation. The research was conducted on the basis of permissions of Provincial Nature Conservation Authorities and the Minister of Environment (DOPog A-2/03/al.; DOPog A-6/04/al.; DLOPiK-op/Ozgi-4200/IV.D-16/6568/06/aj.) and thanks to courtesy of the management of Moravsky kras Landscape Protection Park (Chráněná krajinná oblast Moravský kras). 29

30 5.2. Places of research The most important criteria of choice of places of research was among other things large number of particular bat populations. Data of refugioclimate were collected in the years in the following underground systems of Poland and the Czech Republic (Table 2): Table no. 2. List of underground systems in which research was conducted. Name of underground system longitude latitude Sewers and shopfloor in Police E 14 32' 5 N 53 33' 4 Międzyrzecz Fortified Region E 15 29' 2 N 52 23' 4 Mopkowy Tunel (Barbastelle Tunnel) E 15 12' 4 N 51 48' 4 Cold store in Cieszków E 17 22' 3 N 51 37' 1 A cellar in Ładza E 17 87' 1 N 50 84' 7 Szachownica Cave E 18 48' 2 N 51 31' 5 Underground systems of Tarnowskie Góry and Bytom E 18 49' 5 N 50 24' 8 Fortifications in Nysa E 17 18' 5 N 50 29' 4 A drift in Sławniowice E 17 16' 4 N 50 20' 1 Drifts in forest administration region of Senderki E 23 34' 2 N 50 32' 2 The Balcarka Cave E 16 45'2 N 49 22' 3 Sloup Šošůvka Cave E 16 44'1 N 49 24' 3 30

31 Sewers and shopfloor in Police Network of underground sewers 4000 m long and shopfloor which are remains of prewar factory of aviation fuel (synthetic petrol) Hydrier Werke Politz. The biggest winter habitat of bats in Western Pomerania (780 specimens winter period 2003). Six bat species winter here: Barbastella barbastellus, Myotis myotis, Myotis brandtii, Myotis daubentonii, Myotis nattereri, Plecotus auritus. It is located in Zachodniopomorskie province. The facility has been included into the Natura 2000 network the purpose of which is to preserve specific types of habitats and species which are considered valuable and endangered in entire Europe (area code: PLH320015). Międzyrzecz Fortified Region Refugium Nietoperek encompasses a vast network of old underground fortifications i.e. 30 km of reinforced concrete bunkers, m under ground level. They form a part of so called Międzyrzecz Fortified Region constructed by the Nazi in the years The underground system is connected with the surface by several vertical ventilation shafts and corridors leading to the bunkers. The area encompasses the most important winter habitat of bats in Central Europe and their feeding grounds. About 30 thousand bats winter here (Kokurewicz et al., 2013). The most numerous are: Myotis daubentoni, Myotis myotis, Plecotus auritus and Myotis nattereri. The following species also occur: Barbastella barbastellus, Myotis dasycneme, Myotis bechsteinii, Eptesicus serotinus, Myotis brandtii, Myotis mystacinus, Pipistrellus pipistrellus, Plecotus austriacus. It is located in Lubuskie province. The facility has been included into the Natura 2000 network the purpose of which is to preserve specific types of habitats and species which are considered valuable and endangered in entire Europe (area code: PLH080003). Mopkowy Tunel (Barbastelle Tunnel) Underground drain of a former factory located near Krzystkowice with the outlet to the Bóbr river. The largest known in Poland winter grouping of Barbastella barbastellus. Around 2000 specimens winter here. Several specimens of Myotis daubentonii and Plecotus auritus also occur here. It is located in Lubuskie province. The facility has been included into the Natura 2000 network the purpose of which is to preserve specific types of habitats and species which are considered valuable and endangered in entire Europe (area code: PLH080024). 31

32 Cold store in Cieszków It is a spacious underground brick ice cellar located in a wood in the vicinity of the village of Cieszków. Probably it was built in the 19 th century and served then to the needs of the palace in Cieszków. Later, it was used as a cold store by various users. It is located in Dolnośląskie province. It is one of the largest winter habitats of bats in Poland. The guide Natura 2000 ( ) lists 200 specimens of Barbastella barbastellus, Myotis myotis and Eptesicus serotinus, Myotis daubentonii, Myotis nattereri, Plecotus auritus, Plecotus austriacus. Apart from playing the role of hibernaculum, the cold store is also important for bats which migrate. Late in summer and autumn a large number of specimens are observed which were not spotted here in winter period, for which it is a mating site. It is now secured by Forest District Office in Milicz. A number of changes was introduced in the structure of the facility the purpose of which was to improve microclimatic conditions to make them optimal for bats. Among other things additional partition walls were constructed as well as a pool which ensures appropriate humidity. The facility has been included into the Natura 2000 network the purpose of which is to preserve specific types of habitats and species which are considered valuable and endangered in entire Europe (area code: PLH020001). A cellar in Ładza A small cellar in a building of a former school (currently management of Stobrawski Landscape Protection Park) in Ładza in Opolskie province. Each year 2 specimens of the following species winter here: Plecotus auritus and Plecotus austriacus. It is located in Opolskie province. Szachownica Cave Proglacial cave system in Upper Jurassic limestone in the central part of Wieluńska Upland. The cave system is composed of five separate caves separated by excavation of the quarry. They used to form one cave, destroyed during exploitation of limestone performed by local inhabitants till Currently, the place is treated as one cave system of total length of 1000 m, which before destruction was probably longer than 2 km. It is one of the largest winter habitats of bats in Poland. Each year over 1000 bats representing 10 species hibernate in the cave: Barbastella barbastellus, Myotis dasycneme, Myotis bechsteinii, Myotis myotis, Eptesicus serotinus, Myotis brandtii, Myotis daubentonii, Myotis mystacinus, Myotis nattereri, Plecotus auritus. 32

33 It is located in Śląskie province. The facility has been included into the Natura 2000 network the purpose of which is to preserve specific types of habitats and species which are considered valuable and endangered in entire Europe (area code: PLH240004). Underground systems of Tarnowskie Góry and Bytom Underground excavations remaining after exploitation of heavy metals ore. One of the largest underground systems in the world. The excavations were formed from the 12 th to the 20 th century. Currently they cover over 300 km of corridors as well as numerous chambers and pits. The underground system encompasses 5 drain adits, numerous shafts and outcrops in quarries. It is probably second largest winter habitat of bats in Poland. The following 10 species of bats were observed here to winter: Myotis myotis, Myotis nattereri, Myotis mystacinus, Myotis brandtii, Myotis daubentonii, Myotis bechsteini, Myotis emarginatus, Eptesicus serotinus, Plecotus auritus, Plecotus austriacus (Kłys, 2008). It is located in Śląskie province. The facility has been included into the Natura 2000 network the purpose of which is to preserve specific types of habitats and species which are considered valuable and endangered in entire Europe (area code: PLH240003). Fortifications in Nysa A vast defensive complex of buildings from the 19 th century with a large number of corridors, constructed in topographic low of the valley of Nysa Kłodzka river, currently located in the town park in Nysa. One of the most important wintering habitats of bats in Silesia (Hebda 2001). The guide Natura 2000 informs about wintering of the following species: Rhinolophus hipposideros, Barbastella barbastellus, Myotis emarginatus, Myotis bechsteinii, Myotis myotis. It is located in Opolskie province. The facility has been included into the Natura 2000 network the purpose of which is to preserve specific types of habitats and species which are considered valuable and endangered in entire Europe (area code: PLH160001). A drift in Sławniowice Underground corridor located near the village of Sławniowice on the premises of a marble quarry. About 200 specimens of lesser horseshoe bat Rhinolophus hipposideros winter here. It is the largest known winter habitat of lesser horseshoe bat in the Polish part of Sudety mountains (Kepel et al., 2005). In the vicinity there is a breeding colony of this species. It is located in Opolskie province. The facility has been included into the Natura 2000 network the purpose of which is to preserve specific types of habitats and species which are considered valuable and endangered in entire Europe (area code: PLH160004). 33

34 Drifts in forest administration region of Senderki They are drifts remaining after exploitation of sandstone for production of milling stones in Forestry Commission of Zwierzyniec and on private grounds west from a village of Potok Senderki, behind a grove intersected by a road which is a continuation of the main road which leads through the aforementioned village. The drifts are located at the bottom of tree-covered ravines which cut into fields. It is one of the most interesting winter colonies of bats in Lublin region. Nine bat species winter here: Barbastella barbastellus, Myotis dasycneme, Myotis bechsteinii, Myotis myotis, Myotis brandtii, Myotis daubentonii, Myotis mystacinus, Myotis nattereri, Plecotus auritus. It is located in Lubelskie province. The facility has been included into the Natura 2000 network the purpose of which is to preserve specific types of habitats and species which are considered valuable and endangered in entire Europe (area code: PLH060020). The Balcarka Cave (Jeskyně Balcarka) Balcarka Caves are located in a valley close to a small town of Ostrov u Macochy. Underground maze of corridors, crevices and chambers is created on two levels. The cave is also a valuable paleontological and archaeological site. Bones of Pleistocene animals, instruments made of bone and stone and a bonfire of people from the Old Stone Age were found there. Subsequent parts of the cave were successively discovered in the years and are characterized by rich, various and colourful dripstones. It is one of the most important wintering places of Rhinolophus hipposideros in Europe. It is located in southern Moravia in the Czech Republic. The facility has been included into the Natura 2000 network the purpose of which is to preserve specific types of habitats and species which are considered valuable and endangered in entire Europe (area code: CZ Moravský kras). Sloup Šošůvka Cave (Sloupsko-šošůvské jeskyně) An extensive complex of chambers, corridors and underground chasms created on two levels. Abundant cave fauna (bears, lions, hyenas...) was found there. Remains of Neanderthal man from 120,000 years ago were also found in that place. It is one of the most important wintering places of Rhinolophus hipposideros in Europe (Zima et al., 1994). It is located in southern Moravia in the Czech Republic. The facility has been included into the Natura 2000 network the purpose of which is to preserve specific types 34

35 of habitats and species which are considered valuable and endangered in entire Europe (area code: CZ Moravský kras) General overview of devices and methodology of measurement of hibernation environment (refugioclimate) So far microclimatic (ecoclimatic) research concerned mostly general state of atmosphere of an underground system and rarely microclimate of refuges (Kłys et al., 2005; Kłys, Wołoszyn, 2005; Kłys, 2008), which are direct places of wintering of bats. Microclimate of refuges many times differs greatly from microclimatic background of the underground system. One may use various measuring devices, but methodology of measurement should be precisely described. So far it was impossible to introduce to measuring technology a device which would measure a sum of factors having influence on hibernation comfort of a bat. Most of works concerning microclimatic conditions in a place of occurrence of bats lack the description of methodology of measurement and kind of devices used, which makes correct interpretation of such data impossible (Kłys et al., 2005). Usually influence of the measuring person on the result of measurement is not taken into account as well as time for stabilization of the device, which is often 30 minutes or even longer (Caputa, Kłys, 2005). Apart from measurement of ecoclimate of a refuge the background of the underground system was also measured. Physical factors which determine microclimate of underground systems are discussed below Air temperature (T) The measure of empirical temperature is usually a change in volume or pressure of a standard body which is in the state of thermodynamic balance with the body the temperature of which is measured. There are theoretical and empirical scales of temperatures. The first group includes e.g. scale of perfect gas, thermodynamic scale of temperatures. Empirical scales based on empirical data includes International Practical Temperature Scale. Depending on the way of heat transfer between the sensor and the body the temperature of which is determined; devices are divided into contacting ones called 35

36 thermometers, non-contacting ones (pyrometers) and special ones. In research on bats (estivation, hibernation) the range of measurements should be between C and 50 0 C. A frequent measuring device used so far in measuring temperature was Assman aspiration psychrometer. However, values obtained in this method bear major errors and these measurements also average the values (drawing in larger quantities of air). They give only a very general picture of microclimatic values and they are totally useless to measure refugioclimate. In the literature there is a notion of dry-bulb temperature (T s ) - it is temperature displayed by a normal (i.e. dry) thermometer and if there is no reference to the kind of temperature, it means that dry-bulb temperature is concerned. Wet-bulb temperature (T w ) is the temperature shown by a wet thermometer, e.g. in a psychrometer or covered by ice. It should be pointed out that in small underground systems and relatively stable microclimate, measurements performed with the use of traditional methods are hindered and bear grave errors, also due to presence of the researcher and people who accompany him or her, as well as time of their stay in the place where the measurement is performed (Fig. 4). There is a number of publications concerning influence of tourism in caves on their microclimate, including temperature (Kwiatkowski, Piasecki, 1989; Piasecki, 1996, 1996a; Pflitsch et. al., 1999; Zelinka, 2002; Piasecki et. al., 2007). Measurement of temperature and air flow in a refuge should be performed from the side the air comes in (Fig. 5). For the purpose of this research a thermoanemometer and a gas parameter gauge made by SENSOTRON, specially modified and calibrated to the needs of recording, were used. Special attention was paid to graduate thermometers (devices) in relation to a bench-mark in various ranges of temperature before the measurements were performed. In the immediate place of hibernation of bats temperature was measured with the use of an extension arm (aluminium rod), in such a way, that the observer did not interfere with readings of the gauge, always against the stream of inflowing air. 36

37 Fig. 4. Influence of human presence on measurement of temperature (9 Mar 2002). Measured at the place bat hibernation. Vertical lines show the range of data omitted in analyses (Kłys 2003). 37

38 Substrate temperature (T ch - temperature of the surface of hibernation) To measure temperature of a side wall of an excavation or surface of objects which are close to wintering bats, thermometers or non-contacting devices are used. Usually temperature is measured with thermoelectric or resistance thermometers, less often expansion thermometers are utilized. For the purpose of this work an electric contact thermometer made by SENSOTRON was used. In order to enlarge the surface of contact silicone of high thermal conductivity was utilized Air flow velocity (v) A very important element of ecoclimate which should be registered is measurement of air flow velocity, both the background of ecoclimate and directly in the place of hibernation. This parameter should be approached very carefully. In case of the background of an underground system these measurements require determining average velocity in time and certain cross section or determining spot speed for refugioclimate. In order to measure spot and average velocity in time anemometers, impact pressure tubes, flowmeters, hot-wire anemometers and katathermometers are used. Air flow velocity in underground systems is usually given in m/s, m/min and sometimes cm/s (100 cm/s = 1 m/s = 60 m/min). Sometimes these devices have scales in imperial system units in/s, ft/s, yd/s. (1m/s = in/s = ) Specifying air flow in the hibernation place of a bat with the use of anemometers is difficult due to technical reasons. Above all it refers to measurements (recording) of movement of small velocity which occurs in niches. Measurements of air velocity lower than 0.1 ms -1 are difficult or impossible to perform with the use of regular anemometers. Using mechanical anemometers of various types is not as effective as it was expected due to small inertia of receptors, necessity to overcome internal and external friction, as well as small space of a niche. In the current work a specially modified and calibrated to the needs of recording SENSOTRON hot-wire anemometer was used. In order to minimize influence of a human and a wintering bat measurements were performed from leeward side while standing face front to coming air (Fig. 5).

39 In narrow (low) corridors and chambers of small volume emission of heat (3-4 thousand kcal/24h) and breath of the observer may significantly influence results of observation (Kłys et al., 2005). Fig. 5. Performing measurement of air flow and temperature. The arrow shows direction of air flow. An infrared photograph for the author by P. Rutkowski, Flir Systems. In measurements in cross section of a corridor ( background ) air velocity is not same in all points of the section (Fig. 6). The highest velocities are usually in central parts of a corridor, while the lowest ones are at walls. There are often observed streams of air flowing in and out which flow through the entire inside diameter of the opening in one direction or interchangeably, i.e. in the cross section of the corridor two opposite fluxes of air are moving. 39

40 While measuring background due to gross interferences of flows a crosswise division of cross section of an underground system should be made. In order to avoid errors, measurement of velocity in particular determined places should be performed several times (Pawiński et al., 1995) and an arithmetic mean should be calculated from these measurements. Fig. 6. An exemplary distribution of air velocity in a corridor of an underground system. The place of hibernation of a bat is indicated. (The photograph and the drawing are made by the author). 40

41 Relative humidity of air (Rh) Atmospheric air in underground systems is considered to be a mixture of dry air and water vapour. One should remember, that when temperature of air saturated with vapour drops, part of vapour condenses and mist occurs. As temperature of air saturated with steam increases, state of insatiability occurs (it is shown in a Mollier diagram (Biernacki, 1993). Air humidity (relative and absolute) is a value changeable in time and space. It depends on climate, season of the year, intensity of precipitation and direction of air flow (into or out of a cave). Differences in air humidity of a zone next to an entrance and deeper ones may be significant. Air humidity is an equally complex factor as air temperature. It is formed as a result of moisture incoming from the surface, cooling of air inside, becoming damper in contact with groundwater flows and infiltration water (Kwiatkowski, Piasecki, 1989). To measure humidity the following methods are used: gravimetric, condensation (dew point), psychrometric, hygroscopic ones as well as hygrometer sensors. To measure relative humidity of background of the underground system usually an Assman aspiration psychrometer was used of an accuracy up to 0,2 0 C. Despite very precise measurement of relative humidity the devices used so far (Assman aspiration psychrometer) are totally useless for measuring refugioclimate. The author used an electronic gas parameter gauge made by SENSOTRON. It allowed to measure relative humidity in microniches. The measurement was performed while standing face front to coming air in order to minimize influence of the human and the wintering bat. The bats often use seeps of water from walls (Fig. 7); air humidity is then higher only in the immediate proximity of the wintering bat. 41

42 Fig. 7. In an environment where humidity is lower then the desired one, greater mouseeared bats choose microniches of higher humidity of rock and air (photographed by the author) Thermal conduction of materials (λ) Thermal conductivity, thermal conduction coefficient (marked with the symbol of λ) is one of the most important parameters of substance for heat conduction. In same conditions more heat will flow through a substance of higher thermal conduction coefficient. Thermal conductivity is a quantity characteristic for a substance in a given state of aggregation and its phase. It depends on its chemical composition, structure, porosity, state of aggregation and temperature. The substances which best conduct heat are metals, while gases are the poorest conductors. There are significant difficulties in field measurement of thermal conductivity. 42

43 In the work there were no field measurements of conductivity, only a rough analysis of the substance on which bats wintered. Taking into consideration the type of the substrate approximate values of thermal conductivity were used (Table 3). Table 3. Exemplary approximate values of thermal conductivity of materials which one may encounter at bat hibernation (grey fields show average values of the factor). The unit of the thermal conduction coefficient in SI is J/(m s K) = W m -1 K -1 (watt per meter kelvin). material Thermal conductivity in (W m -1 *K -1 ). The average value is given Air Expanded polystyrene 0,03; 0,06; 0,1 Wood 0,04; 0,12; 0,21 Rubber 0,16 Water 0,5; 0,55; 0,6 Brick 0,6-0,15; 0,6; 0.69; 1.31 Concrete 0,8; 1,0; 1,28 Limestone 1,33 Soil 0,6; 2,3; 4 Sandstone 1,83; 2,4; 2,90 Marble 2,07; 2,5; 2,94 Granite 1,73; 2,8; 3,98 Cast iron 55; Iron 71,8; 80,2; 55.4; 34.6; 60,5 43

44 Air pressure (p) Air pressure depends on basic quantities, which must be taken into consideration in research on underground systems. Knowing the value of pressure is useful for estimation of velocity of flow, volume of flux and mass of air. During comparison of data of refugioclimate with humidity and temperature a program calculating Mollier diagram was used (Wykres i-x Molliera). In underground systems depending on the way of measurement the following items are used: devices to measure absolute pressure: mercury barometers, aneroid barometers, barolux, micro-barolux; devices to measure pressure above or below atmospheric: micromanometers, manometers, differential manometers. According to the principle of operation one may distinguish: liquid gauges, elastic pressure gauges and electric converters. The author used an electronic gas parameter gauge made by SENSOTRON, which had a built-in barometer. In the current work values of pressure of refugioclimate were converted into values for 1000 hpa of absolute pressure (the program Wykres i-x Molliera). It facilitates comparison of data of relative humidity from different measuring points as well as days of measurement Level of cooling (K a ) While entering the state of hibernation a body of a bat may give up heat to the environment by radiation, vaporization, convection and conduction. The amount of heat which is given up in convection depends on thermal conductivity of the body of the bat and difference of temperatures of skin and air or rock that surrounds the body. In certain combination of such factors as: temperature, air movement and humidity one may assume that the bat achieves comfort of hibernation. In order to determine optimal values for hibernation of bats dry-bulb temperature, relative humidity and air flow velocity was measured in the place of hibernation. Thermal comfort is determined by measurement of intensity of cooling with the use of wet Hill s katathermometer. The quantity of cooling power of the atmosphere, which is intensity of cooling K a, is expressed by loss of heat from 1 cm 2 of surface in 1 second (Frycz, 1974). 44

45 The unit of intensity of cooling is 1 Kata degree mcal/cm 2 s. Due to the commonly binding SI the intensity of cooling effect of the atmosphere should be expressed in W/m 2. The unit of intensity of cooling is NK a (new Kata degree) expressed in W/m -2. Due to frequent use of the old unit in literature, a conversion formula has been given. NK a = 42 x K a [W/m -2 ] There are the following empirical relations between the cooling effect of the atmosphere given in Kata degrees and air velocity υ and its temperature T expressed in 0 C (Budryk, 1961). υ 1ms -1 Where: w air velocity, ms -1 T w wet-bulb temperature K w = (0,35 + 0,85 ³ w) (36,5 -T w ) Due to measuring difficulties (in our case low temperature, disturbances by human presence itself, heating up the katathermometer, repeating the measurement at least 5 times and above all else difficulty in placing it close to the hibernating bat) the above mentioned formula was used. So the following components were measured: air flow velocity and temperature in the proximity of a hibernating bat, but the wet-bulb Hill s katathermometer was not used due to the above mentioned measuring errors. Unfortunately, there is no direct formula which allows to calculate wet-bulb temperature Tm on the basis of dry-bulb temperature T s and relative humidity Rh. Due to the fact that dry-bulb temperature T s and relative humidity Rh was measured, not wet-bulb temperature T w which is necessary for the formula, T w was calculated with the use of the computer program Wykres i-x Molliera the purpose of which is to simplify calculations connected with transformations of humid air. The list of devices used during research as well as their parameters are included in Appendix no

46 46

47 6. Wintering strategies While describing microclimatic conditions in hibernation place of bats, the sum of basic physical parameters usually is not usually taken into consideration, the methodology of measurement is not described (so is it not known what was measured in fact??), it is also usually forgotten to present wintering strategy of bats. The works are not numerous (Zukal et al., 2005). We do not know if the bats winter individually, socially, are they hidden in crevices. Therefore in the current work the following division (depending on participation of physical factors which have influence on cooling of a bat) into wintering strategies is proposed (Fig. 8): Individual wintering Ia hanging freely Ib hanging on the wall Ic in a crevice Group wintering (social) IIa hanging freely IIb hanging on the wall IIc in a crevice In social behaviour we may distinguish further complexity e.g. in strategy IIb particular specimens may winter placed loosely one next to the other, overlapping or create a cluster in which particular specimens are placed one on another (Fig. 9). 47

48 Fig. 8. Depending on contact with physical factors wintering strategies used by the bats taking into account factors (air flow, humidity, rock temperature, air temperature) and possibility to give up heat by convection or conduction. 48

49 Fig. 9. Social behaviour in IIb strategy. a - Myotis myotis - overlapping, b - Barbastella barbastellus - cluster (one on another), c - Myotis daubentonii and Myotis nattereri - loosely one next to another (Photographed by the author). Of course not all species use all strategies and they are not observed in equal percentage. It is commonly known, that lesser horseshoe bat always uses Ia strategy and no other behaviour during hibernation was observed. Brown long-eared bat is often observed in strategy Ia ; Ib rarely in Ic, though own observations while catching them alive (Kłys, 2008) show, that the last strategy is probably more common, but difficult to observe. They are also found in mixed colonies with other species. 49

50 50

51 7. Thermal comfort of a hibernating bat The balance of exchange of matter and energy between a hibernating specimen and environment, meaning general qualitative and quantitative metabolism must include: sum of heat transmitted to the system from environment sum of heat transmitted outside by the system thermal effect of processes occurring inside the system If the loss of energy in the organism is balanced by reserve substances, the organism of a hibernating bat is in the state of thermal (energy) equilibrium. The organism of a hibernating bat has possibility to achieve thermal balance in quite wide range of physical parameters of surrounding environment and their changes thanks to the system of thermoregulation of the organism. It has to maintain continuously controlled temperature, a bit higher from ambient, but within the optimal range (which we do not know usually). The parameters which characterize metabolism in quantitative terms in the state of energy balance depend, similarly as in humans, from constant factors (species, size, weight, sex, age of the specimen) and variable ones (regulation of heat exchange of the body in contact with environment). The energy effect of metabolism may be referred to body weight or to surface area of the organism, or, for best results, to both factors simultaneously. The organism of a bat may change surface area of the body (by spreading or folding wings, hiding or sticking out ears (Plecotus), snuggling to objects or protruding from them). It is then relatively labile (flexible). Only highly specialized species like e.g. Rhinolophus hipposideros have different strategy of managing physical conditions of environment. A bat wrapped in his patagium can part it or wrap it more tightly, which gives him possibility to achieve thermal balance and ensure comfort of hibernation (Fig. 10). 51

52 Fig. 10. Possibility to regulate influence of physical factors on the organism by Rhinolophus hipposideros (Photographed by the author). The climate of wintering place (refugioclimate) are numerical values of those physical and chemical parameters which have influence on amount of heat exchanged between the organism of a hibernating bat and its environment. The amount of heat transmitted from an organism to environment is called heat loss. In order to characterize the hibernation environment a notion of refugioclimate is used, which means the state of climatic conditions occurring in natural way or created artificially in small space surrounding a hibernating bat, hence formed among other things by influence of: - average air temperature, - amount of moisture in the air, - air flow velocity, - temperature of surrounding objects. 52

53 Common (joint) influence of these factors on the organism of a hibernating bat may be called thermal comfort of hibernation, which should be presented in numerical form; that would be the hibernation coefficient. An attempt was made here to integrate these values into wet Kata cooling power degree (see chapter 3.2). These values are only an approximation of the searched hibernation coefficient and are not noticeable enough in all examined species. So by the thermal comfort of a wintering bat one should conceive such a state of satisfaction of a specimen (group) in thermal conditions of environment in which it feels neither warmth nor coolness. The necessary condition to feel thermal comfort is achieving the state of thermal balance of organism. Such a state is characterized by levelling the amount of heat of metabolism with energy exchanged between the organism of a hibernating bat and environment (Fig. 11). Fig. 11. Possibilities of regulation of thermal condition of a bat s organism. A hibernating bat may stay in various conditions characterized by various values of physical quantities enumerated above. The most beneficial refugioclimate is created by such conditions in which a bat feels well and the heat management of its organism is most economical. This state is achieved in various mutual combinations of temperature, relative humidity and air flow velocity and it is called the state of thermal comfort. Due to biological differences in particular population there is no possibility for all specimens of a given species which are present in a place of a given refugioclimate to feel well and comfortably in given thermal conditions. For this reason it should be assumed that the optimal refugioclimate is a state in which a possibly high proportion (e.g %) of a hibernating bat population accepts prevailing physical conditions. Sometimes there may be a disturbance of the process of carrying heat from organism to the environment. Such a state may occur when production of heat of metabolism is higher than the amount of heat that may be carried to the environment in given conditions. Such a state is called discomfort of hibernation. 53

54 Bats may be forced to change the hibernation place; it happens for two reasons: When the organism of a hibernating bat transmits heat too slowly, its body temperature rises. Whereas it transmits too much heat it cools down below the optimal value. The organism dies or must use more energy to get warm (loses reserves faster) or fly away to find favourable conditions to survive. In both cases a bat tries to change its hibernation place. An organism transmits heat to the environment thanks to existence of difference of temperatures of a specimen and environment and difference in saturated vapour pressure in temperatures of skin and environment. While entering the state of hibernation the body of a bat may give up heat to the environment by: radiation, vaporization, convection and conduction (Fig. 12). Fig. 12. Processes of heat exchange between an organism and the environment. 54

55 7. 1. Conduction Heat transfer from an organism to the environment through conduction may occur when a part of the body is in contact with solid body, such as a side wall, a recess, a rock crevice, etc. Depending on a strategy of hibernation adopted by a bat (see chapter 5.1. Wintering strategies) and surface area of contact of the body of the animal with substrate, amount of heat transferred this way will be very varied. Currently, due to the scope of research only approximate values may be discussed (1 70%?) Radiation In bats which hibernate on the surface, outside underground systems, in heat exchange between their organisms and surrounding environment a crucial role may be played by solar radiation and thermal radiation originating on Earth. In underground systems only thermal radiation occurs. Each surface emits flux of radiation of energy depending on its absolute temperature and emissivity according to Stefan-Boltzmann law Convection Removal of heat through convection from a body which is not covered by fur is characterized by coefficient of thermal insulation of a layer of hair at the wall, air around the body, which is inverse of coefficient of heat transfer. Value of thermal insulation is connected with surface layer of air around a body and constitutes a crucial element of total thermal resistance of the organism. In case of high velocity of cool air flow quantity and quality of hair plays an important role. The amount of heat which is given up in convection depends on thermal conductivity of the body of the bat and difference of temperatures of skin and air (rock) that surrounds the body. In normal conditions an organism loses most heat through vaporization (Schmidt-Nielsen, 1997). Relative humidity of air surrounding a bat has enormous significance in this case. When humidity is close to saturated condition giving up excess heat from an organism to environment may be hindered or even impossible. Through convection, which is direct lifting from skin surface, intensity of heat exchange rises on the surface of body itself. Giving up heat from an organism through 55

56 convection and conduction depends on air velocity. Mechanisms which regulate amount of heat created in an organism must operate in such a way that in short time thermal balance becomes even, otherwise it may be impossible to maintain body temperature at a constant level Vaporization Influence of relative humidity is often more important than nutritional status of animals (Thomas, Cloutier, 1992; Thomas, 1995). Even in conditions of high relative humidity (90-98%) the loss of moisture may reach significant values. The loss of moisture occurs through external surface of the animal s body and through breathing. Flux of heat is always directed from warmer to colder body, in underground conditions it means that as long as air or rock that surrounds a bat is cooler than its skin, heat will be transmitted from skin surface to air or rock. Through convection, which is direct lifting or conduction of heat from skin surface, intensity of heat exchange rises on the surface of the body of a bat itself. It is the only method of decreasing body temperature in a short time. Slow entering into hibernation state (long-term decrease in body temperature) would cause an increase in energy loss. The quantity of cooling power of the atmosphere, which is intensity of cooling, expresses loss of heat from 1 cm 2 of surface in 1 second. So a bat needs to shorten the time of entering into the state of hibernation as much as it is possible. Therefore, the factor of wind (air flow velocity) has two tasks; first, to shorten the time of entering into the state of hibernation, second, to carry away heat. Giving up heat from an organism through convection and conduction depends on air velocity and is in direct proportion to the difference of temperatures of air and body surface. During winter in the temperate zone bats enter into a state of deep torpor. During that period body temperature drops within the range of 1-2 o C above ambient (Hock, 1951; Henshaw and Folk, 1966; Herried and Schmidt Nielsen, 1966; McNab, 1974). At such difference in temperature costs of metabolism during hibernation are exceptionally low in comparison to euthermic animals (Hock, 1951; Herried and Schmidt-Nielsen, 1966; Thomas et al., 1990a, 1990b). Wintering in this period depends on fat reserves, which are source of energy during hibernation, and effective usage of them. After all, hibernation period may last even 6 months. Assuming then, that the difference between the temperature of a hibernating bat and the temperature of the refugioclimate is 1 C, the cooling factor of the atmosphere will be of value within tenth of Kata degree (at the same parameters of atmosphere). It depends, of course, also on thermal conductivity of a bat s body. In typical hibernation temperature drops almost to ambient, energy loss will be then even lower, which allows to survive several months without feeding, and energy to sustain life is drawn from fat reserves. 56

57 So far no clear-cut methodology was found to assess in thermal terms the environment in which humans live and work as well as bats hibernate. In adequate number of points one may measure certain changeable parameters of this environment, which shape the state of heat exchange between an organism and its environment. Estimation if a refugioclimate is proper constitutes a necessary condition to work out indicators of thermal comfort. Engineers, psychologists and physiologists are interested in this issue from their point of view. Current work constitutes then certain possibilities to predict and evaluate given physical conditions of refugioclimate from the point of view of hibernation. Conditions of environment should be considered as comfortable if a hibernating bat feels neither warmth nor cold. 57

58 58

59 8. Choice of place of hibernation Introduction Research on bats is methodologically complicated due to their specific lifestyle. Information about bats is often gathered as a result of an accidental encounter; rarely it is due to planned long-term research. Earlier research undertook estimation of changes in number of bats in relation to microclimate of the underground system. The majority of research concerned thermopreferendum of bats (Gaisler, 1970; Bauerova and Zima, 1988). Harmata, (1969) stated that temperature is the most important factor responsible for hibernation. We know that each species has specific needs as far as hiding place is concerned in relation to types of shelter, temperature, humidity and stability of environment (Kunz, Anthony, 1982). According to the author the above-mentioned conditions are not sufficient parameters to describe refugioclimate. (Kłys et al., 2002; Kłys, 2004, 2008) suggests and points to one more factor, which is air flow velocity and in this paper also thermal conduction of rocks and strategy of hibernation. In order to evaluate influence of the sum of these factors on the choice of hibernation place the author performed measurements of refugioclimate of dry-bulb temperature (T s - n = 6389), relative humidity (Rh - n = 6389), air flow (v - n = 6389) and atmospheric air pressure (p - n = 6389). Only part of results of physical values of refugioclimate of chosen six species of bats were used for the analysis. It produced in total items of data. The analysis did not take into consideration data from mixed social strategies (multi-species groups) and species which were observed sporadically during research (pond bat Myotis dasycneme; Brandt s bat Myotis brandtii; whiskered bat Myotis mystacinus; Geoffroy s bat Myotis emarginatus; Bechstein s bat Myotis bechsteinii; serotine bat Eptesicus serotinus; and grey long-eared bat Plecotus austriacus). Majority of data published so far which concerned physical quantities (T s, v and Rh) from hibernation places of bats are difficult to interpret and sometimes simply unacceptable. In the published works the detailed methodology of measurements is not usually given and one does not know if the described data refer to hibernation place (refugioclimate). For example Nagy, Postawa (2011) informs that a measurement was performed 1.5 m below hibernation place. It is unacceptable because physical parameters in underground systems often differ significantly even in short distances (several 59

60 cm from a hibernating bat). Using a katathermometer for measuring air flow velocity by Kokurewicz (2004) should be excluded absolutely from analysis. Using Assman psychrometer by the above-mentioned author is also methodologically erroneous (see chapter 5.3. General overview...). Data collected by the current author also contain certain error. Usually we do not know in which phase of hibernation a given specimen was in the moment of measurement of physical quantities of refugioclimate. The full range of physical conditions of refugioclimate was not always available. Therefore, mainly at small number of data the formulae of mathematical functions of a hibernation place will be subject to changes along with supply of new data on physical quantities from hibernation places. However, the conditions of research on refugioclimate were given for potential verification and supplementation of data Whether there is a relation of choosing by bats higher temperature at first rather than in later months of hibernation In the scientific literature temperature (T = T s ) is usually taken into consideration without other microclimatic factors. It is also mentioned that bats choose higher T at the beginning of hibernation than in the latter months; it is probably due to the fact that small underground systems have higher internal temperature in autumn since they are heated after summer. It is more probable that bats are very economic and choose the most optimal places as far as heat loss is concerned. Due to the above mentioned facts a comparison was performed of measured T s of refugioclimate for Western Barbastelle Barbastella barbastellus in three months: December, January, February (APPENDIX Table 1). The analysis of statistical significance was performed with the use of Kruskal Wallis test since the groups, as far as specific strategies are concerned, differ in size and some of them are low in number, e.g. there were 29 measurements in December, so influence of abnormality of factorization on results induces non-parametric analyses. Particular months of hibernation were compared with the use of Dunn s test (APPENDIX Table 2). Highly significant differences (p<0.0001) of dry-bulb temperature between months were observed. Highly significant differences concern comparison of December with February (p=0.0009) and January with February (p<0.0001). Whereas the difference between December and January is not significant (p=1.0000). It means highly significant increase of dry-bulb temperature in the hibernation place in February in comparison with December and January (Fig. 13). 60

61 Fig. 13. Results of measurements of T s in the hibernation place of Barbastella barbastellus during three months. The performed analyses show errors in previous thinking about choices of bats concerning hibernation places Joint or separate analysis of physical factors of refugioclimate In earlier publications concerning hibernation of bats one may observe lack of comprehensive view on the aforementioned physical factors. As a general rule, authors limit themselves to providing only one of them, usually temperature, less often temperature and humidity. 61

62 As part of research on refugioclimate the present author performed measurements of T s, Rh and v. The author suggests that none of these factors should be examined separately, but jointly. Due to this fact the author analysed dry-bulb temperature, relative humidity and air flow velocity in the view of correlation between the examined features and choice of those features by wintering bats (Fig. 14, 15, 16) and eigenvalues were established (APPENDIX Table 3). Fig. 14. Correlation diagrams of dispersion of dry-bulb temperature and relative humidity. Barbastella barbastellus place of hibernation 62

63 Fig. 15. Correlation diagrams of dispersion of dry-bulb temperature and air flow velocity. Barbastella barbastellus place of hibernation. Fig. 16. Correlation diagrams of dispersion of relative humidity and air flow velocity. Barbastella barbastellus place of hibernation. 63

64 It was observed that the component corresponding to the first largest eigenvalue explains only 39.5% of total variance. Second component explains 31.7% of total variance and the third one explains 28.8% which sums up to 100% of observed variance. Even first two components jointly explain only 71.2% of observed variance which means that to explain adequately the observed variation all three components are necessary which in turn means lack of more efficient reduction of dimensions. It means that the description requires direct use of all three examined parameters (T s ; Rh; v). On one hand it is very difficult to simplify these three parameters into one. On the other hand it means that each of the parameters of microclimate has got very large influence, so it is not legitimate to limit oneself just to T s Comparison of physical factors (T s ; Rh; v) of refugioclimate of chosen bat species Therefore, six chosen bat species were compared regarding T s dry-bulb temperature, Rh relative humidity and v air flow velocity. In order to compare these data of refugioclimate basic statistics have been presented (APPENDIX Table 5). Data were subjected to multiple analysis with Tukey s test. The performed Tukey s multiple comparison test for unequal numbers has shown significant differences between mean values of T s ; Rh and v for particular bat species (APPENDIX Table 6). Single Factor Analysis of Variance has also shown that numbers are so big that the assessed parameter, which is the mean, is near to real one independently from departures from normal distribution. Taking T s into consideration highly significant differences (p<0.0001) between species were observed. Four groups of wintering bats can be distinguished that significantly differ from one another, that is: the lowest dry-bulb temperature concerned PAR and the highest concerned MYN; the species RHH and BAR are characterized by similar dry-bulb temperature and likewise the species MYD and MYM are characterized by similar dry-bulb temperature. Therefore, the sequence is as follows (from the lowest to the highest): PAR, RHH with BAR, MYD with MYM, MYN. Taking Rh into consideration highly significant differences (p<0.0001) between species were observed. Four groups can be distinguished that significantly differ from one another, that is: the lowest relative humidity concerned BAR and the highest concerned RHH; the species PAR and MYD are characterized by similar relative humidity and likewise the species MYN and MYM are characterized by similar relative humidity. Therefore, the sequence is as follows (from the lowest to the highest): BAR, PAR with MYD, MYM with MYN, RHH. 64

65 Taking v into consideration highly significant differences (p<0.0001) between species were observed. The species RHH and MYN occurred at the lowest air flow and were the species which significantly diverged from the remaining ones, but they also differed between themselves and so the lowest air flow was observed in case of RHH. Among other species PAR was in the significantly lower air flow than MYD, while the species BAR and MYM did not differ significantly from each other nor from the two above mentioned ones taking the medium values. Therefore, the sequence is as follows (from the lowest to the highest): RHH, MYN, PAR, BAR with MYM, MYD. Results of the performed analysis have been illustrated with box-and-whisker plots, which are drawn up on the basis of values of descriptive statistics and which show distribution of ordered values of features (Fig. 17, 18, 19). Ryc. 17. Diversification of chosen bat species in relation to the choice of dry-bulb temperature (T s ). 65

66 Fig. 18. Diversification of chosen bat species in relation to the choice of relative humidity (Rh). Fig. 19. Diversification of chosen bat species in relation to the choice of air flow velocity (v). 66

67 Due to great difficulty in simplifying these three parameters into one parameter wet Kata cooling power degrees were used, the values of which were compared between species (APPENDIX Table 7). Taking (K w ) into consideration highly significant differences (p<0.0001) between species were observed. One may distinguish three groups which differ significantly from one another, which means that their lowest value concerned RHH and MYN. The MYM, MYD and BAR species are characterized by K w which is not different significantly and likewise BAR and PAR species are characterized by similar K w. Therefore, the sequence is as follows (from the lowest to the highest): RHH with MYN, MYM with MYD with BAR with PAR (Fig. 20). The table with precise Tukey s multiple comparison tests is attached in the appendix (APPENDIX Table 8). Single Factor Analysis of Variance shows that numbers are so big that the assessed parameter, which is the mean, is near to real one independently from departures from normal distribution. Fig. 20. The scope of the preferred range of values of wet Kata cooling power degrees by the chosen species of bats. 67

68 As the result of the analysis one has to observe that each of the species has highly specific preferendum of hibernation defined by variables T s, Rh and v. There are species for which some of the values of parameters of refugioclimate are similar, but they differ in relation to another of the enumerated parameters Analysis of particular species Authors of earlier works did not take into consideration strategies of wintering in their descriptions of microclimatic conditions in hibernation places of particular species of bats. Whether they winter separately, socially; if they are e.g. hidden in crevices (see chapter 6, Wintering strategies). Particular species, divided into hibernation strategies, are presented below Lesser Horseshoe Bat Rhinolophus hipposideros (Bechstein, 1800) Diversification of chosen physical quantities (T s ) dry-bulb temperature, (Rh) relative humidity and (v) air flow velocity is presented in (APPENDIX Table 9) and Fig. 17, 18, 19. Lesser horseshoe bat hibernates in underground systems using only the strategy of hanging freely ( Ia ). In its winter habitat it does not create colonies (Wołoszyn, 2001). The animals during hibernation are usually at a distance of centimetres from one another. The species prefers wintering places of high air humidity, it often amounts to 100% (Harmata, 1994), which is confirmed by the author s studies. Hibernation temperature fluctuates from 2 C to 14 C (Burbank and Young, 1934; Kowalski, 1953; Daan and Wichers, 1968; Gaisler, 1970; Harmata, 1987 Roer and Schober, 2001; Paksuz et al., 2007). The author observes in the result of his research that at lower values of T s and higher velocity v the species chooses higher relative humidity Rh which is justified for energy reasons. At higher dry-bulb temperature T s and lower air flow velocity, relative humidity is lower. It should be remembered that the biggest number of specimens wintered at T s between 7,0 0 C and 8,8 0 C, at relative humidity Rh % and air flow velocity v 0,04 0,09 m/s and within these ranges one should see optimal conditions of hibernation. 68

69 Due to great difficulty in simplifying these three parameters into one parameter wet Kata cooling power degrees were used, the values of which were compared between species (Fig. 20). The value of K w is presented in figure 21. Fig. 21. The scope of values of wet Kata cooling power degrees chosen by lesser horseshoe bat Rhinolophus hipposideros. Empirical diagram and discriminant of a quadratic equation were determined where if v and T s are known, Rh can be calculated (Fig. 22, 23). A polynomial function of the second degree was matched to the points in the diagram of 3D dispersion. 69

70 Fig. 22. Empirical diagram of dispersion of three variables T s, Rh and v for Ia hibernation strategy of lesser horseshoe bat Rhinolophus hiposideros. Figures in squares mark the value of K w. In the blue field interquartile range of the observed specimens has been marked. RHH "Ia" Fig. 23. Empirical diagram of dispersion of three variables Ts, Rh and v for Ia hibernation strategy of lesser horseshoe bat Rhinolophus hiposideros. 70

71 Rh = 78, ,1184 T + 316,644 v - 0,1567 T T - 14,2066 T v ,271 v v where: v means air flow velocity m/s T s - means dry-bulb air temperature in C Rh means relative humidity in % At lower values of T s and higher velocity v RHH chooses higher relative humidity which is justified for energy reasons. At higher dry-bulb temperature T s and lower air flow velocity, relative humidity is lower. It should be remembered that the biggest number of specimens winters at T s between 7 0 C and 8,8 0 C, at relative humidity Rh % and air flow velocity v 0,04 0,09 m/s and within these ranges one should see optimal conditions of hibernation. In the dispersion diagram of three variables T s, Rh and v for lesser horseshoe bat Rhinolophus hipposideros in the range of lower and upper quartile of variables T s, Rh and v the diagram adopts linear function described by the formula below. Rh = 97,7708 0,0456 T + 27,2202 v The species worked out a specific style of hibernation Ia of hanging freely at the roof of an underground system and wrapping its patagium around itself. As far as refugioclimate conditions are concerned, the species requires stable microclimate of the interior of an underground system (Zukal et al., 2005) Western Barbastelle Barbastella barbastellus (Schreber, 1774) Diversification of chosen physical quantities depending on hibernation strategy has been presented in Table 10 (APPENDIX) and Fig. 24, 25, 26. This species chooses for hibernation dry and cold places, with hibernation temperature -3,0-6,5 (Gais1er, 1970; Bogdanowicz, 1983; Bogdanowicz and Urbanczyk, 1983; Lesinski, 1986); it can endure short-lived drops in temperature to -9 C. It tolerates humidity from 60 to 90%. These values have been given despite the lack of precisely described methodology of measurement in the cited literature. The species is very flexible, during hibernation almost all basic strategies of hibernation were observed. Only IIa strategy was not noted, it should be expected to be observed during further research. It proves very high flexibility as far as the choice of hibernaculum is concerned. The most frequently observed strategy was strategy Ib, in which specimens wintered on concrete (λ 1) and rubber (λ 0.16). These strategies 71

72 are the most frequently spotted ones, however, one should expect that specimens hidden in crevices may be equally or even more frequent than those observed. The performed Tukey s multiple comparison test for unequal numbers has shown significant differences between mean values of T s ; Rh and v for particular strategies of BAR. Detailed Tukey s multiple comparison test between particular hibernation strategies is presented in Table 11 (APPENDIX). In case of T s the highest one was noted in strategy Ic which was not different only in comparison to strategy Ia. Strategies Ib concrete and Ib rubber differed between each other highly significantly (p=0.0010) in relation to dry-bulb temperature, taking into consideration that the lower one concerned rubber. In case of relative humidity strategies Ib concrete and Ib rubber differed between each other highly significantly (p = 0,0082) and the higher relative humidity concerned concrete. Relative humidity of strategy IIc did not differ significantly from all other ones. In case of air flow strategies Ib concrete and Ib rubber differed between each other highly significantly (p = 0,0001) and the higher air flow concerned rubber. Air flow of strategy IIc did not differ significantly from all other ones. In order to interpret gathered parameters easier, diversification of chosen strategies has been presented in relation to T s, Rh and v with the use of box-and-whisker plots (Fig. 24, 25, 26). Fig. 24. Diversification of chosen hibernation strategies of Western Barbastelle Barbastella barbastellus in relation to choice of (T s ) dry-bulb temperature in 0 C. 72

73 Fig. 25. Diversification of chosen hibernation strategies of Western Barbastelle Barbastella barbastellus in relation to choice of (Rh) relative humidity in %. Fig. 26. Diversification of chosen hibernation strategies of Western Barbastelle Barbastella barbastellus in relation to choice of (v) air flow velocity in m/s. 73

74 Due to great difficulty in simplifying these three parameters into one parameter wet Kata cooling power degrees were used, the values of which were compared between strategies (Table 12 APPENDIX). The analysis of significance was performed with the use of Kruskal Wallis test since the groups, as far as specific strategies are concerned, differ in size and some of them are low in number, so influence of abnormality of factorization on results induces non-parametric analyses. Detailed multiple comparison with the use of Dunn s test (APPENDIX Table 13) between strategies of choice of wet Kata cooling power degrees allowed to observe highly significant (p < 0,0001) differences between factorization of results of wet Kata cooling power degrees depending on strategy. Strategies Ib concrete and Ib rubber differed from each other highly significantly (p < 0,0001) in relation to Kata degrees and the lower one concerns hibernation on concrete (λ 1.0 W m -1 K -1 ), (Fig. 27). Fig. 27. Diversification of chosen strategies of Barbastella barbastellus in relation to choice of wet Kata cooling power degrees. 74

75 Strategy Ia, though the sample was very small (n=6), does not differ significantly from other strategies in dry-bulb temperature (T s ) of refugioclimate and it seems that specimens wintering in such manner base on the same parameters of refugioclimate as when using other strategies. Wet Kata cooling power degrees, which connect the described parameters of refugioclimate with one another, diversify strategy Ia. Significant statistical differences are observed between Ib (λ 1) and IIb. However, when we take a look at humidity (Rh), statistically it differs significantly from Ic and IIb, while in case of air flow velocity it differs from Ib wintering on concrete (λ 1) and IIb. It doesn t significantly differ statistically from strategy Ib of those wintering on rubber (λ 0.16), which is a perfect isolator. So cooling will depend above all else on convection both in strategy Ia and Ib (on rubber). Wet Kata cooling power degrees also do not show statistically significant differences between these strategies. Strategy Ib (λ 1) differs in relation to wet Kata cooling power degrees from all strategies except IIb and IIc, but when components are analysed (T s ; Rh; v) it differs statistically significantly from these two strategies with dry-bulb temperature. Lower T s of strategy IIb (λ 1) than Ib (λ 1) causes gathering into groups. Strategy Ib (λ 0.16) differs in relation to wet Kata cooling power degrees from Ib (λ 1); IIb. Analysing components (T s ; Rh; v) it differs statistically significantly in relation to T s from Ib (λ 1); Ic in relation to Rh Ib (λ 1); Ic and IIb. In relation to v differs only from IIb. Strategy Ic differs in relation to wet Kata cooling power degrees from Ib (λ 1); IIb. Analysing components (T s ; Rh; v) it differs statistically significantly in relation to T s from all except Ia, in relation to Rh from Ia, Ib (λ 1) and Ib (λ 0,16). It differs in relation to v from Ib (λ 1) and IIb. Strategy IIb differs in relation to wet Kata cooling power degrees from all except Ib (λ 1). Analysing components (T s ; Rh; v) it differs statistically significantly in relation to T s from Ib (λ 1); Ic, in relation to Rh from Ia ; Ib (λ 0,16). It differs in relation to v from Ia ; Ib (λ 0,16) and Ic. Strategy IIc differs in relation to wet Kata cooling power degrees from Ib (λ 1) and IIb. Analysing components (T s ; Rh; v) it differs statistically significantly in relation to T s from Ic, in relation to Rh they do not differ from each other statistically. In relation to v they do not differ statistically. 75

76 Therefore, choice of strategy depends not only on value of wet Kata cooling power degrees (K w ), but the change may be also caused by an increase or decrease in one of the components T s ; Rh or v and also it may depend on thermal conductivity of substrate (λ). Therefore, it has been concluded that conditions of refugioclimate cause BAR to choose the most appropriate strategy of hibernation to adapt to existing conditions of physical factors. Empirical diagrams and discriminants of a quadratic equation were determined for majority of strategies (for strategy Ic and IIc due to the fact that the group was small no empirical discriminant of quadratic equation could be determined) where at known air flow velocity v and dry-bulb temperature T s one may calculate value of relative humidity Rh (Fig. 28, 29, 30, 31, 32, 33, 34 35, 36). A polynomial function of the second degree was matched to the points in the diagram of 3D dispersion. 76

77 Strategy Ia Rh = 394, ,7881 T + 19,6503 v + 2,3965 T T + 47,5522 T v 411,4992 v v. Fig. 28. Empirical diagram of dispersion of three variables T s, Rh and v for Ia hibernation strategy of Western Barbastelle Barbastella barbastellus. Figures in squares mark the value of K w. In the blue field interquartile range of the observed specimens has been marked Fig. 29. Empirical diagram of dispersion of three variables T s, Rh and v for Ia hibernation strategy of Western Barbastelle Barbastella barbastellus. In the dispersion diagram of three variables T s, Rh and v for Barbastella barbastellus in the range of lower and upper quartile of variables T s, Rh and v the diagram adopts linear function described by the formula below. Rh = 79, ,0833 T - 30,1263 v A polynomial function of the second degree was matched to the points in the diagram of 3D dispersion. 77

78 Strategy Ib concrete Rh = 110,5609-5,2788 T - 52,2461 v + 0,141 T T + 7,8808 T v + 12,9543 v v Fig. 30. Empirical diagram of dispersion of three variables T s, Rh and v for Ib hibernation strategy of Western Barbastelle Barbastella barbastellus. Figures in squares mark the value of K w. In the blue field interquartile range of the observed specimens has been marked. Fig. 31. Empirical diagram of dispersion of three variables T s, Rh and v for Ib concrete hibernation strategy of Western Barbastelle Barbastella barbastellus. In the dispersion diagram of three variables T s, Rh and v for Barbastella barbastellus in the range of lower and upper quartile of the variables the diagram adopts linear function described by the formula below. Rh = 91,3035-1,7913 T v 78

79 Strategy Ib rubber Rh = 61, ,4684 T + 51,4689 v - 0,0018 T T - 7,0409 T v + 9,6291 v v Fig. 32. Empirical diagram of dispersion of three variables T s, Rh and v for Ib hibernation strategy of Western Barbastelle Barbastella barbastellus. Figures in squares mark the value of K w. In the blue field interquartile range of the observed specimens has been marked. Fig. 33. Empirical diagram of dispersion of three variables T s, Rh and v for Ib rubber hibernation strategy of Western Barbastelle Barbastella barbastellus. In the dispersion diagram of three variables T s, Rh and v for Barbastella barbastellus in the range of lower and upper quartile of the variables the diagram adopts linear function described by the formula below. For strategy Ic, IIb the number of data is too small and for IIa there is no data at all. Rh = 78,8284-0,7532 T + 0,7608 v 79

80 Strategy Rh = 51, ,846 T + 68,2567 v - 0,3917 T T - 9,2499 T v + 0,4008 v v Fig. 34. Empirical diagram of dispersion of three variables T s, Rh and v for IIb hibernation strategy of Western Barbastelle Barbastella barbastellus. Figures in squares mark the value of K w. In the blue field interquartile range of the observed specimens has been marked. In the dispersion diagram of three variables T s, Rh and v for Barbastella barbastellus in the range of lower and upper quartile of the variables the diagram adopts linear function described by the formula below. 80 Rh = 95,8774-2,1269 T + 2,7887 v Fig. 35. Empirical diagram of dispersion of three variables T s, Rh and v for IIb hibernation strategy of Western Barbastelle Barbastella barbastellus.

81 Fig. 36. Diagram of points of three variables T s, Rh and v for IIc hibernation strategy of Western Barbastelle Barbastella barbastellus Daubenton s Bat Myotis daubentonii (Kuhl, 1817) Diversification of chosen physical quantities (T s ) dry-bulb temperature, (Rh) relative humidity and (v) air flow velocity in relation to strategy of hibernation is presented in Table 14 (APPENDIX) and Fig. 37, 38, 39. The analysis of significance was performed with the use of Kruskal Wallis test since the groups, as far as specific strategies are concerned, differ in size and some of them are low in number, so influence of abnormality of factorization on results induces non-parametric analyses. This species chooses for hibernation underground systems of hibernation temperature of (Haagen and Arnold, 1955; Daan and Wichers, 1968; Gaisler, 1970; Daan, 1973; Bogdanowicz, 1983, 1994; Bogdanowicz and Urbanczyk, 1983; Lesinski, 1986; Harmata, 1987; Mazing, 1987; Corbet and Harris, 1991; Macdonald & Barrett, 1993; Bogdanowicz, 1994; Kłys et al., 2002). 81

82 Relative humidity % (Bogdanowicz and Urbariczyk, 1983; Kłys et al., 2002; Kokurewicz 2004). Air flow velocity in the underground system is said to be 0,04 ms -1 (Kokurewicz 2004). In artificial caves in the Netherlands air flow reached from 0,1 to 0,4 ms -1 (Daan, 1973), while in the Nietoperek underground fortification system with many entrances it reaches from 0,0 to 3,1 ms -1 (Kokurewicz, 1999). Values of physical factors have been given despite the lack of precisely described methodology of measurement. Detailed multiple comparison between strategies was performed with the use of Dunn s test (Table 15 APPENDIX). In case of dry-bulb temperature one highly significant difference was observed (p = 0,0037) between Ib and IIb, which may suggest that at lower T s and same other parameters Daubenton s bats start to group. Colonies observed during research were very loosely positioned one next to another Fig. 9.c. Whereas in case of relative humidity and air flow no significant differences were observed (p > 0,05). Fig. 37. Diversification of chosen hibernation strategies of Daubenton s bat Myotis daubentonii in relation to choice of (T s ) dry-bulb temperature in 0 C. 82

83 Fig. 38. Diversification of chosen hibernation strategies of Daubenton s bat Myotis daubentonii in relation to choice of (Rh) relative humidity in %. Fig. 39. Diversification of chosen hibernation strategies of Daubenton s bat Myotis daubentonii in relation to choice of (v) air flow velocity in m/s. 83

84 Due to great difficulty in simplifying these three parameters into one parameter K w were used, the values of which were compared between strategies (Table 16 APPENDIX, Fig. 40). The analysis of significance was performed with the use of Kruskal Wallis test since the groups, as far as specific strategies are concerned, differ in size and some of them are low in number, so influence of abnormality of factorization on results induces non-parametric analyses. Detailed multiple comparison between choice of strategy and K w was performed with the use of Dunn s test (Table 17 APPENDIX). No significant differences were observed (p > 0,05) between strategies. Fig. 40. Diversification of chosen strategies of Myotis daubentonii in relation to choice of wet Kata cooling power degrees. 84

85 The species was observed to use strategy Ia (n=1); Ib, Ic (n=3) and IIb. Data of strategy Ia and Ic are so few, that only strategies Ib and IIb were compared with each other. In relation to wet Kata cooling power degrees between the enumerated strategies no significant statistical differences were observed. During an analysis of components (T s ; Rh; v) significant statistical differences were observed in relation to T s. Drop in T s causes gathering into groups. At similar values of Rh and v. Empirical diagrams and discriminants of a quadratic equation were determined for majority of strategies (for strategy Ia and Ic due to the fact that the sample was small no empirical discriminant of quadratic equation could be determined) where at known air flow velocity v and dry-bulb temperature T s one may calculate value of relative humidity Rh (41, 42, 43, 44). A polynomial function of the second degree was matched to the points in the diagram of 3D dispersion. 85

86 Strategy Ib Rh = 118,0028-8,3647 T + 6,559 v + 0,523 T T + 0,0635 T v - 2,2267 v v Fig. 41. Empirical diagram of dispersion of three variables T s, Rh and v for Ib hibernation strategy of Daubenton s bat Myotis daubentonii. Figures in squares mark the value of wet Kata cooling power degrees. In the blue field interquartile range of the observed specimens has been marked. Fig. 42. Empirical diagram of dispersion of three variables T s, Rh and v for Ib hibernation strategy of Daubenton s bat Myotis daubentonii. In the dispersion diagram of three variables T s, Rh and v for Myotis daubentonii in the range of lower and upper quartile of variables T s, Rh and v the diagram adopts linear function described by the formula below. 86 Rh = 77, ,9333 x + 4,9312 y

87 Strategy IIb Rh = 95,713-2,0012 T - 3,3507 v + 0,1161 T T + 0,666 T v + 0,2346 v v Fig. 43. Empirical diagram of dispersion of three variables T s, Rh and v for IIb hibernation strategy of Daubenton s bat Myotis daubentonii. Figures in squares mark the value of K w. In the blue field interquartile range of the observed specimens has been marked. Rh % In the dispersion diagram of three variables T s, Rh and v for Myotis daubentonii in the range of lower and upper quartile of variables T s, Rh and v the diagram adopts linear function described by the formula below. Rh = 85, ,172 x + 2,791 y Fig. 44. Empirical diagram of dispersion of three variables T s, Rh and v for IIb hibernation strategy of Daubenton s bat Myotis daubentonii. 87

88 Natterer s Bat Myotis nattereri (Kuhl, 1817) Diversification of chosen physical quantities (T s ) dry-bulb temperature, (Rh) relative humidity and (v) air flow velocity in relation to strategy of hibernation is presented in Table 18 (APPENDIX) and Fig. 45, 46, 47. This species chooses for hibernation underground systems of hibernation temperature of ((Daan and Wichers, 1968; Gais1er, 1970; Bogdanowicz, 1983; Bogdanowicz and Urbanczyk, 1983; Lesinski, 1986). Hibernaculum may be performed in two ways. In places where underground wind blows and temperature is around 6, it falls into winter sleep in deep crevices, often with other species, inter alia Daubenton s bat. Whereas if air flow is low, temperature fluctuates between 9 and 11, then Natterer s bat winters on the walls of corridors (Konkurewicz et al., 1996). Values of physical factors have been given despite the lack of precisely described methodology of measurement. Diversification of Natterer s bat Myotis nattereri in relation to (T s ) dry-bulb temperature, (Rh) relative humidity and (v) air flow velocity depending on strategy of hibernation is presented in Table 19 and Fig. 45, 46, 47. The analysis of significance was performed with the use of Student s t-test (we have two groups of high numerical strength, so the estimated means are close to real) Table 19 (APPENDIX). In case of T s highly significant (p < 0,0001) differences between both strategies were observed. Lower T s concerned strategy IIb. In case of Rh highly significant (p = 0,0238) differences between both strategies were observed. Lower relative humidity concerned strategy IIb. In case of v not significant (p = 0,2467) difference between both strategies was observed. 88

89 Fig. 45. Diversification of chosen hibernation strategies of Natterer s bat Myotis nattereri in relation to choice of (T s ) dry-bulb temperature in 0 C. Fig. 46. Diversification of chosen hibernation strategies of Natterer s bat Myotis nattereri in relation to choice of (Rh) relative humidity in %. 89

90 Fig. 47. Diversification of chosen hibernation strategies of Natterer s bat Myotis nattereri in relation to choice of (v) air flow velocity in m/s. Due to great difficulty in simplifying these three parameters into one parameter wet Kata cooling power degrees were used, the values of which were compared between strategies (Table 19 APPENDIX). Fig. 48. The analysis of significance was performed with the use of Student s t-test (we have two groups of high numerical strength, so the estimated means are close to real) (Table 19 - APPENDIX). No significant difference was observed (p < 0,9590) 90

91 Fig. 48. Diversification of chosen strategies of Myotis nattereri in relation to choice of wet Kata cooling power degrees. The species was observed to use strategies Ib and IIb. In relation to wet Kata cooling power degrees between the enumerated strategies no significant statistical differences were observed. During an analysis of components (T s ; Rh; v) significant statistical differences were observed in relation to T s. Statistically significant differences were observed also in values of Rh and v. Lower humidity and flow were observed at strategy IIb. Drop in T s causes gathering into groups. Empirical diagrams and discriminants of a quadratic equation were determined for majority of strategies (for strategy Ic and IIc due to the fact that the sample was small no empirical discriminant of quadratic equation could be determined) where at known air flow velocity v and dry-bulb temperature T s one may calculate value of relative humidity Rh (49, 50, 51, 52). A polynomial function of the second degree was matched to the points in the diagram of 3D dispersion. 91

92 Strategy Ib Rh = 153, ,7292 T - 30,6339 v + 0,7278 T T + 3,8154 * T v - 2,6111 v v Fig. 49. Empirical diagram of dispersion of three variables T s, Rh and v for Ia hibernation strategy of Natterer s bat Myotis nattereri. Figures in squares mark the value of K w. In the blue field interquartile range of the observed specimens has been marked M YN "Ib" Fig. 50. Empirical diagram of dispersion of three variables T s, Rh and v for Ia hibernation strategy of Natterer s bat Myotis nattereri. In the dispersion diagram of three variables T s, Rh and v for Myotis nattereri in the range of lower and upper quartile of the variables the diagram adopts linear function described by the formula below. Rh = 87, ,3219 * T+0,5162 v 92

93 Strategy IIb Rh = 85,5426-0,2928 T + 53,9946 v + 0,0638 T T - 4,1104 T v-19,7282 v v Fig. 51. Empirical diagram of dispersion of three variables T s, Rh and v for Ia hibernation strategy of Natterer s bat Myotis nattereri. Figures in squares mark the value of K w. In the blue field interquartile range of the observed specimens has been marked. M Y N "IIb" Fig. 52. Empirical diagram of dispersion of three variables T s, Rh and v for Ia hibernation strategy of Natterer s bat Myotis nattereri. In the dispersion diagram of three variables T s, Rh and v for Myotis nattereri in the range of lower and upper quartile of variables T s, Rh and v the diagram adopts linear function described by the formula below. Rh = 83, ,618 T + 3,994 v 93

94 Greater Mouse-eared bat Myotis myotis (Borkhausen, 1797) Diversification of chosen physical quantities (T s ) dry-bulb temperature, (Rh) relative humidity and (v) air flow velocity in relation to strategy of hibernation is presented in Table 20 (APPENDIX) and Fig. 53, 54, 55. The analysis of significance was performed with the use of Kruskal Wallis test since the groups, as far as specific strategies are concerned, differ in size and some of them are low in number, so influence of abnormality of factorization on results induces non-parametric analyses. Winter shelters of this species were noticed both in caves and artificial underground systems. It winters individually or in numerous clusters which run into several hundred specimens; it chooses places for hibernation with temperature of -4,0-12,0 (Haagen and Arnold, 1955; Daan, Wicheres, 1968; Gaisler, 1970; Bogdanowicz, 1983; Bogdanowicz, Bagrowska-Urbańczyk, Urbańczyk, 1983; Lesiński,1986; Harmata, 1987; Webb et al., 1996). Values of physical factors have been given despite the lack of precisely described methodology of measurement. Detailed multiple comparison with the use of Dunn s test between the choice of strategy of greater mouse-eared bat is presented in Table 21 (APPENDIX). In case of dry-bulb temperature the distinctly significantly lowest one was observed in case of Ib dolomite and at the same time in this strategy the highest, significantly higher from other, relative humidity. In case of air flow only strategy IIb differed significantly from a few others having the lowest value of air velocity. In order to interpret gathered parameters easier, diversification of chosen strategies has been presented in relation to T s, Rh and v with the use of box-and-whisker plots (Fig. 53, 54, 55). 94

95 Fig. 53. Diversification of chosen hibernation strategies of greater mouse-eared bat Myotis myotis in relation to choice of (T s ) dry-bulb temperature in 0 C. Adjacent value Outliers Adjacent value Outliers concrete dolomite Rh % Fig. 54. Diversification of chosen hibernation strategies of greater mouse-eared bat Myotis myotis in relation to choice of (Rh) relative humidity in %. 95

96 Fig. 55. Diversification of chosen hibernation strategies of greater mouse-eared bat Myotis myotis in relation to choice of (v) air flow velocity in m/s. Due to great difficulty in simplifying these three parameters into one parameter wet Kata cooling power degrees were used, the values of which were compared between strategies (Table 22 APPENDIX, Fig. 56). The analysis of significance was performed with the use of Kruskal Wallis test since the groups, as far as specific strategies are concerned, differ in size and some of them are low in number, so influence of abnormality of factorization on results induces non-parametric analyses. In Table no. 23 (APPENDIX) the choice between strategies was analysed through detailed multiple comparisons with the use of Dunn s test. Generally, the group which differs from some other groups is IIb Fig. 56. Diversification of chosen strategies of Myotis myotis in relation to choice of K w.

97 The species was observed to use strategy Ia ; Ib (λ 1); Ib (λ 1.33) and IIb. Strategy IIb was divided in terms of numerical strength. IIb (2-20); IIb (20-40); IIb (40-80); IIb (^80) In relation to K w significant statistical differences were observed between strategies Ib (λ 1) and IIb (40-80). Significant statistical differences were found also between strategy IIb (20-40); strategy IIb (40-80) and IIb (^80). Analysing Rh significant statistical differences were observed between Ib and IIb (40-80) as well as between IIb (2-20) and IIb (40-80). In relation to v significant statistical differences were observed between Ib (λ 1) and strategy IIb (40-80) as well as between IIb (2-20) and IIb (40-80) and also between IIb (20-40) and IIb (40-80). Comparison of social groupings to individual specimens. It may result from a fact that the number of specimens in relation to total number of examined specimens was below 1% of all registered specimens. They may be accidental specimens which do not fully hibernate. Out of the examined species this one is the most difficult to interpret. This fact may be caused by e.g. influence of family groupings which are not fully connected with microclimate. Difficulties with attributing this species to particular physical factors are mentioned by Zukal et al., (2005). Empirical diagrams and discriminants of a quadratic equation were determined for majority of strategies, where at known air flow velocity v and dry-bulb temperature T s one may calculate value of relative humidity Rh (Fig. 57, 58, 59, 60). A polynomial function of the second degree was matched to the points in the diagram of 3D dispersion. 97

98 Strategy Ib Rh = 120,7662-7,6628 T - 0,9409 v + 0,4365 T T + 0,681 T v - 2,1618 v v Rh % Fig. 57. Empirical diagram of dispersion of three variables T s, Rh and v for Ib hibernation strategy of greater mouse-eared bat Myotis myotis. Figures in squares mark the value of K w. In the blue field interquartile range of the observed specimens has been marked. In the dispersion diagram of three variables T s, Rh and v for Myotis myotis in the range of lower and upper quartile the diagram adopts linear function described by the formula below. Rh = 83, ,43 x + 3,3719 y 98 Fig. 58. Empirical diagram of dispersion of three variables T s, Rh and v for Ib hibernation strategy of greater mouse-eared bat Myotis myotis with an indicated area of the most frequently observed (25-75%).

99 Strategy IIb Rh = 100,2182-2,9481 T + 9,0412 v + 0,1734 T T + 0,1071 T v - 5,839 v v Rh % Fig. 59. Empirical diagram of dispersion of three variables T s, Rh and v for Ia hibernation strategy of greater mouse-eared bat Myotis myotis. Figures in squares mark the value of K w. In the blue field interquartile range of the observed specimens has been marked. Fig. 60. Empirical diagram of dispersion of three variables T s, Rh and v for IIb hibernation strategy of greater mouse-eared bat Myotis myotis. In the dispersion diagram of three variables T s, Rh and v for Myotis myotis in the range of lower and upper quartile the diagram adopts linear function described by the formula: Rh = 89,6914-0,1217 x + 3,9555 y 99

100 Brown Long-eared Bat Plecotus auritus (Linnaeus, 1758) Diversification of chosen physical quantities (T s ) dry-bulb temperature, (Rh) relative humidity and (v) air flow velocity in relation to strategy of hibernation is presented in Table 24 (APPENDIX) and Fig. 60, 61, 62. The analysis of significance was performed with the use of Kruskal Wallis test since the groups, as far as specific strategies are concerned, differ in size and some of them are low in number, so influence of abnormality of factorization on results induces non-parametric analyses. This species chooses for hibernation underground systems with hibernation temperature of 0,0-9,0 (Daan, Wichers, 1968; Gaisler, 1970; Bogdanowicz, 1983; Bogdanowicz, Bagrowska-Urbańczyk, Urbańczyk, 1983; Lesiński, 1986; Harmata, 1994; Webb et al., 1996; Kłys, Wołoszyn, 2005). Values of physical factors have been given despite the lack of precisely described methodology of measurement. Detailed multiple comparisons with the use of Dunn s test were performed - Table 25 (APPENDIX). In case of dry-bulb temperature a significant (p=0.0116) difference was observed between strategies Ia and Ib ; the lower temperature concerns strategy Ia. In case of relative humidity a highly significant (p=0.0059) difference was observed between strategies Ia and Ib ; the lower relative humidity concerns strategy Ib. No significant difference was observed in air flow velocity in relation to strategy. In order to interpret gathered parameters easier, diversification of chosen strategies has been presented in relation to T s, Rh and v with the use of box-and-whisker plots (Fig , 63.). Fig. 61. Diversification of chosen hibernation strategies of brown long-eared bat Plecotus auritus in relation to choice of (T s ) dry-bulb temperature in 0 C. 100

101 Fig. 62. Diversification of chosen hibernation strategies of brown long-eared bat Plecotus auritus in relation to choice of (Rh) relative humidity in %. Fig. 63. Diversification of chosen hibernation strategies of brown long-eared bat Plecotus auritus in relation to choice of (v) air flow velocity in m/s. 101

102 Due to great difficulty in simplifying these three parameters into one parameter wet Kata cooling power degrees were used, the values of which were compared between strategies (Table 26 APPENDIX, Fig. 64). The analysis of significance was performed with the use of Kruskal Wallis test since the groups, as far as specific strategies are concerned, differ in size and some of them are low in number, so influence of abnormality of factorization on results induces non-parametric analyses. Detailed multiple comparisons with the use of Dunn s test were also performed - Table 27 (APPENDIX). No significant difference was observed in wet Kata cooling power degrees in relation to strategy. Fig. 64. Diversification of chosen strategies of Plecotus auritus in relation to choice of K w. K w Min-max 102

103 The species was observed to use strategies Ia ; Ib and Ic. In relation to wet Kata cooling power degrees between the strategies no significant statistical differences were observed. During an analysis of components (T s ; Rh; v) significant statistical differences were observed in relation to T s, and Rh between strategies Ia and Ib. Strategy Ic was very rare. Lower T s and higher Rh causes brown long-eared bats to choose strategy Ia and higher T s and lower Rh causes them to choose strategy Ib. In various types of underground systems the real number of hibernating brown long-eared bats is probably higher than the registered one, since they hide in narrow and inaccessible for an observer crevices (Bogdanowicz, Urbańczyk, 1983). Own observations in Underground systems of Tarnowskie Góry and Bytom confirm such observations. Empirical diagrams and discriminants of a quadratic equation were determined for majority of strategies, where at known air flow velocity v and dry-bulb temperature T s one may calculate value of relative humidity Rh (Fig. 65). A polynomial function of the second degree was matched to the points in the diagram of 3D dispersion. 103

104 Strategy Ia Rh = 94, ,7873 T - 11,6063 v + 0,0657 T T-7,9423 T v + 135,1346 v v Rh % Fig. 65. Empirical diagram of dispersion of three variables T s, Rh and v for Ia hibernation strategy of lesser horseshoe bat Plecotus auritus. Figures in squares mark the value of K w. In the blue field interquartile range of the observed specimens has been marked. In the dispersion diagram of three variables T s, Rh and v for Plecotus auritus in the range of lower and upper quartile T s, Rh and v the diagram adopts linear function described by the formula: Rh = 96, ,7776 x - 32,1041 y Fig. 66. Empirical diagram of dispersion of three variables T s, Rh and v for Ia hibernation strategy of brown long-eared bat Plecotus auritus. 104

105 Strategy Ib Rh = 105,4839-1,7793 T - 41,8804 v - 0,1091 T T + 5,5733 T v + 1,3347 v v Rh % Fig. 67. Empirical diagram of dispersion of three variables T s, Rh and v for Ib hibernation strategy of brown long-eared bat Plecotus auritus. In the dispersion diagram of three variables T s, Rh and v for Plecotus auritus in the range of lower and upper quartile T s, Rh and v the diagram adopts linear function described by the formula: Rh = 104,9037-2,5695 x + 0,6854 y PAR "Ib" Fig. 68. Empirical diagram of dispersion of three variables T s, Rh and v for Ib hibernation strategy of brown long-eared bat Plecotus auritus. 105

106 9. Summary Hibernation is a precisely regulated hypometabolic process and a hypothermal state of an organism, not passive submission to influence of the environment (Janicki, Cygan-Szczegielniak, 2006). In order to fight against coolness bats are able to choose a place of hibernation and an option of hibernation. Despite the fact that the rule of loss of energy through conduction, convection, radiation and vaporization is widely known (Cena et al., 1986), still works appear which discuss only one of the factors, mainly T a (ambient temperature), which attempt to determine on this basis hibernation comfort, reasons of grouping of specimens an other relations. Sometimes the data are supplemented by relative humidity (Rh). As it was proved (chapter 8) the use of at least three examined parameters (T s ; Rh; v) is required for description of hibernation place. Of course the substrate on which bats winter (λ degree of thermal conductivity) has very important influence on the choice of hibernation strategy. It is also obvious that at the same temperature (T a ) and variable other discussed factors the thermal comfort (of a bat and a human as well) is different and even discomfort occurs (Evola and Popov, 2006; Stamou and Katsiris 2006; Heiselberg et al., 2008; Bohojło and Kołodziejczyk 2009). Before winter bats can increase their body mass (m b ) by 30% (Krzanowski, 1961; Kunz et al., 1998). In order to facilitate accumulation of fat, at lower temperatures (T a ) bats can actively change thermal preferences of choice of shelter (Speakman and Rowland, 1999). Only proper management of accumulated reserves leads to survival of unfavourable period which is winter in cool and temperate climate. Proper choice of refugioclimate and strategy in available physical conditions is one of factors which condition survival (Kłys, Wołoszyn, 2010) Air temperature In countries of temperate climate bats and other animals encounter temperatures much lower than 0 0 C. However, body temperature of warm-blooded animals, even in hibernation state, is always above zero (Poczopko, 1990). So far, air temperature was the basic parameter which characterises winter habitats of bats. Harmata (1969) stated that it is the most important factor responsible for hibernation. Thomas et al., (1990) suggest, that ambient temperature is the strongest factor which determines wintering. Bats in natural conditions search actively for optimal temperature for hibernation in order to minimize loss of fat (Kokurewicz, 2004). Temperature differs significantly within and 106

107 between species of bats (Boratyński et al., 2012). It is observed (Kokurewicz, 2004; Boyles et al., 2007; Wojciechowski et al., 2007) that preferences for temperature during hibernation is not a stable phenomenon, but it depends on many internal and external factors as well as energy reserves, availability of favourable thermal conditions. It should be emphasized that not sole temperature, but a set of physical factors is significant, and temperature is only one of several physical components which provide thermal comfort during hibernation (Cena et al., 1986). A view that bats while falling into hibernation state choose places of relatively low temperature from -17 to ca. +15 still persists in the literature (Harmata, 1969, 1973; Nagel, Nagel, 1991). Webb et al., (1996) collected data concerning 34 species of bats which winter in the temperate zone. Bats were found at temperatures between -10 and +21 o C. The present author used for the current publication measurements in hibernation place (refugium or refuge) (n = 5838) specimens of six species of bats which hibernated in the scope 0,7-12 o C. As it was mentioned above it is hard to refer to most of these data since authors provide methodology of measurement with low accuracy and probably air temperature which was registered referred to the state of air of microclimate of the interior, not necessarily refugioclimate, which, as the research of the present author shows, often significantly differ between one another. Statements about hibernation of bats found in literature probably result from the fact that measurements of temperature concerned microclimate of underground systems, not values of refugioclimate. The present author observed similar situations where temperature in an underground system was below 0 0 C; then bats wintered deeply in crevices where temperature was significantly higher. Giving up heat occurred then mainly through conduction (warming up cooling down) Relative humidity of air It is generally known that content of water vapour in the air is one of decisive factors which shape microclimatic conditions of underground systems. The air which is completely saturated with water vapour cannot take warmth from a body of a bat through vaporization without simultaneous increase in temperature of the air itself. Despite the fact that bats do not possess sweat glands, skin is permeable to water and water from bodily fluids permeates onto the surface where it vaporizes. The smaller is relative humidity of air the larger may be the share of cooling the body by vaporization. The air which is saturated with water vapour has more difficulties in taking heat from a bat s body through vaporization. The necessary condition is convection which allows to take heat from a hibernating bat. Some authors (Thomas and Cloutier, 1992; Paksuz et al., 2007) state an important role of this factor. Lesiński, (1986) and Bogdanowicz, Urbańczyk, (1983) 107

108 provide data on relative humidity on the level of 75-95%; they suggest that species which endure lower temperatures during hibernation (including brown long-eared bat) are more flexible as far as humidity requirements are concerned than species which prefer higher temperatures. Bernard et al., (1991) provides values of relative humidity during hibernation within the scope of %. During research the present author observed the scope of relative humidity (n = 5838) of refugioclimate of chosen species on the level of 53,6-100%. Some authors claim that influence of this parameter is more important than nutritional status of animals (Thomas, Cloutier, 1992; Thomas, 1995). Even in conditions of high relative humidity (90-98%) the loss of moisture may reach significant values. It is believed that clusters may facilitate saving energy by lowering temperature and loss of water (Studier, 1970; Procter and Studier, 1970; Stapp et al., 1991; Canals, 1998; Brown, 1999; Jefimow et al., 2011; Wojciechowski et al., 2011; Boratyński et al., 2012). Barbastella barbastellus draws attention due to the fact that out of all examined species it chooses the lowest humidity. It simply avoids values above 90% of relative humidity. Increased humidity in case of this species causes gathering specimens into clusters. So in different species this factor may cause different behaviour which is not fully explained. Boratyński et al., (2012) used for his research pressure (compressibility) of saturated water vapour (WVP), a derivative of relative humidity, which is dependent also on temperature and pressure Air flow velocity So far it was assumed that air flow velocity influences negatively on the choice of place and course of hibernation of bats. It was assumed that they choose places with no air flow (Kallen, 1964; Tinkle and Patterson, 1965). Such conclusions were caused probably by technical difficulties with measurement of small velocities of air below 0,1 ms -1. In result of the performed research the present author concluded that this view is not correct. Results (Kłys et al., 2002; Kłys, 2004, 2008) show that species of bats winter in places close to which significant air flow velocities (drafts) occur and if only structure (morphology) of walls is developed enough, they are able to find microniches in which parameters of air flow are optimal both for the species and the strategy of wintering chosen by the bat. Observations of the present author have clearly shown that lack of air flow as well as increased air flow are unfavourable. Whereas small air flow defined for both the species and its strategy is necessary for hibernation. Hence slow movements of air in underground systems have enormous significance in settling these shelters during winter period. Air velocity is so high here that it causes fast entering into hibernation state, yet low enough not to cause quick cooling out of the hibernating bat. Observation shows that air movement has significant influence on the level of cooling thus on distribution of bats in underground systems during hibernation. Richter et al., (1993) examined caves in which entrances were modified in order to 108

109 facilitate access for people. Such modification worsened hibernacula of bats. From caves of warm climate walls which hampered exchange of air were removed. Within ten years number of bats increased from 2,000 to 13,000. It should be remembered that circulation of air of different velocity occurs in almost all underground systems. Spatial distribution of temperature and air humidity is often distorted by changes of direction and velocity of air flow. Daan (1973) informs that air flow in an artificial underground system in the Netherlands spanned from 0,1 to 0,4 ms -1. Kokurewicz, (2004) provides data that in the underground system Sowia Dolina air flow velocity did not exceed 0,04 ms -1. In the underground system Nietoperek air flow velocity (Kokurewicz, 1999) ranged from 0,0 to 3,1 ms -1. However, as it was mentioned before one may not attribute these values to particular physical factors of refugioclimate and the authors themselves do not analyse these values and do not draw specific conclusions. For the purpose of the discussion calculation of average amount of flow (Kokurewicz, 2004) in an underground system becomes pointless. Regardless of all, bats from a mosaic of air flow velocities choose in a given moment the most beneficial values which depend on other physical quantities (T s, Rh). The performed research show that wintering bats can very precisely match values of air movements to values of remaining examined factors. The scope of air flow velocity of refugioclimate of chosen bat species taken into account in the current work (n = 5838) equalled 0,01-1,42 ms Thermal conduction The present author did not find in the literature data which would indicate taking into consideration thermal conductivity of substrate (λ) as a factor having meaning in the choice of hibernation by bats. The author has demonstrated that it is one of the basic factors conditioning choice of strategy of hibernation. A perfect example which illustrates this phenomenon is Barbastella barbastellus for which data of the same strategy were compared and specimens wintered on various substrates (concrete, rubber) (chapter ) Air pressure In the current work values of atmospheric pressure of refugioclimate were converted into values for 1000 hpa of absolute pressure (the program Wykres i-x Molliera ). It facilitated comparison of data of relative humidity from different measuring points as well as days of measurement. Where differences in pressure were often significant. 109

110 9. 6. Wintering strategies Particular physical factors of refugioclimate differentiate bat species and cause that a bat matches wintering strategies to a given situation. Complexity of hibernation strategy of particular species is relatively high. The lesser horseshoe bat Rhinlophus hiposideros may be considered as an exception (chapter 10 Thermal comfort of a hibernating bat ). A horseshoe bat wrapped in its patagium can part it or wrap it more tightly, which gives him possibility to achieve thermal balance and ensure comfort of hibernation. Similar strategy of covering, but only abdominal part, is used by species of the genus Plecotus. The majority of hibernating species has possibility to cling more or less to the surface on which they hibernate. Parting or tightening their patagium. Holding out or folding ears (Plecotus). All this allows them to adapt to the environment. If certain physical quantities are breached, specimens look for next possibilities to fit into crevices or to group. The objective is always to find comfort of hibernation. Which is such a state of satisfaction in thermal conditions of environment in which the bat feels neither warmth nor coolness. The necessary condition to feel thermal comfort is achieving the state of thermal balance of organism. Such a state is characterized by levelling the amount of heat of metabolism with energy exchanged between the organism of a hibernating bat and the environment. Therefore, it should be accepted that the state of thermal balance for a specimen of a particular species will be similar both in case when it winters individually or in a group, both when it hangs freely in the air or winters on the surface or in a crevice. Cuddling up lowers use of energy e.g. in birds (Wojciechowski et al., 2011), but it is in fact looking for thermal comfort, because in a group the values of thermal comfort may also be breached. Which means that either too cool or too warm is bad. Of course if there are possibilities of choice. Increased energy losses will occur when values of warmth are either higher or lower than optimal ones. It should be noted that gathering into clusters aims also at reducing risk from predators and that there may be emotional, family ties. Bats look for places which are the most appropriate due to physical parameters of an underground system and they adapt to conditions found through choice of strategy of hibernation. 110

111 9. 7. Standard of conditions of measurement In order to compare properly refugioclimate in hibernation places a unified standard of conditions of measurement is required. So far it was impossible to introduce to measuring technology a device which would measure a sum of factors having influence on hibernation comfort of a bat. Ecoclimatic research used to concern general state of atmosphere of underground systems, but the state of atmosphere of refuges, the immediate place of wintering of bats, was not examined. Those researchers did not fully realized the nature of the issue. Most of works concerning microclimatic conditions in a place of occurrence of bats (refugioclimate) lack the description of methodology of measurement and kind of devices used, which makes correct interpretation of such data impossible (Kłys et al., 2005). Therefore, the following elements have to be taken into account on the measurement chart of a hibernating bat (APPENDIX measurement chart): external and internal conditions of the place of wintering of bats as well as the species and strategies of hibernation. See chapter 5 (Material and Methods) Synthesis of chosen physical factors The present author has made an attempt at a synthesis of chosen physical factors (T s, Rh, v, λ, p) attributing them to species and strategies of wintering. The combination of these factors causes the general physical and mental state of a hibernating bat to reach its optimum. Due to complexity of thermal environment of mammals (bats, human) there were many suggestions of indicators which take into account the specificity of the surroundings in a form of one number (Wacławik, 2010). Usually (though not always) this indicator has the dimension of temperature. With the use of it thermal sensations experienced by people in a specific environment or thermal state of an organism were assessed. So far no such indicator has been worked out for particular species of bats. The most appropriate one for research on hibernation seemed to be Kata cooling power degrees (see chapter 5.3.). It allowed in a series of cases to simplify three factors into one. However, not in all cases Kata cooling power degrees describe a refuge of a wintering bat sufficiently enough. According to the present author this problem requires further precise research. Hence the areas of dispersion of three variables of a polynomial function of the second degree have been determined. 111

112 The results of research show that the percentage share of cooling by particular components of physical quantities (T s ; Rh; v) at particular strategies will be different. At strategy Ia the share of cooling by convection will be significant in contrast with Ic where the decisive factor carrying off the excess heat will be conduction. However, the state of comfort in both cases will be similar. Indeed as far as dry-bulb temperature (T s ) is concerned highly significant differences between species were observed. Only species RHH and BAR are characterized by dry-bulb temperature which is not significantly different and likewise species MYD and MYM are characterized by similar dry-bulb temperature. While MYD and MYM can be found in mixed colonies in underground systems, one rather cannot count on such an encounter in case of RHH and BAR since they differ between each other too much as far as relative humidity is concerned. Taking relative humidity into consideration highly significant differences between species were observed. The species PAR and MYD are characterized by relative humidity which is not significantly different and likewise the species MYN and MYM are characterized by similar relative humidity. Here in certain ranges of physical quantities both species can be found in mixed colonies. Taking air flow velocity into consideration highly significant differences between species were observed. The species RHH and MYN occurred at the lowest air flow and were the species which significantly diverged from the remaining ones, but they also differed between themselves and so the lowest air flow was observed in case of RHH. Therefore one can assume that even single physical factors of refugioclimate differentiate the majority of the examined species. More complicated situation will occur if particular strategies are included. It should be stated that responsibility for the choice of hibernation places of the chosen bat species falls on values of physical factors (T s ; Rh; v) and thermal conductivity of materials (λ) on which bats winter. Values of these components make a hibernating bat match hibernation strategy to conditions of microclimate found in the interior of the underground system in which the bat chooses a niche described by physical values of refugioclimate. So particular physical factors of refugioclimate differentiate bat species and cause that a bat matches appropriate wintering strategies to a given situation. Each species has its preferendum in slightly different scope of Kata cooling power degrees differentiated additionally by force of sometimes single factor. It should be assumed that each of bat species has its preferendum (area described by an equation) determined by physical factors (T s ; Rh; v and λ) which mutually supplement one another. They are described by wet Kata cooling power degrees, but supplemented by values of particular components. Sometimes just one of the factors may cause change of hibernation strategy. Particular strategies of a species are probably 112

113 equal as far as energy is concerned. Bats, depending on conditions of environment and possibilities, choose appropriate strategy to survive winter period as economically as possible. Conditions of refugioclimate (abiotic factors) cause that a bat chooses optimal for him conditions of hibernation by matching strategies to ensuing situation. It is obvious that there is possibility of lack of appropriate refugioclimates; a bat must then expend more accumulated energy and feels discomfort. The problem of choice of higher temperature at the beginning of hibernation cannot be considered, because only a set of factors (described above) is responsible for comfort of hibernation. Such a potential choice may result from a fact that in autumn a wintering place is slightly warmer than in spring and it is advantageous to choose strategy Ia, because it is the most economical in terms of energy and along with cooling down the interior of the underground system towards spring gathering into clusters occurs (of course it is dependant on the possibility of finding adequate microclimatic refuges) Theoretical and practical meaning The current dissertation deals with research on choice of hibernation places of the above mentioned species and its relation with conditions of refugioclimate. While undertaking this research practical meaning was also taken into consideration. The obtained results and conclusions from the current paper allow to propose two important theoretical and practical meanings in ecology and protection of bats in various ecosystems. They are: Comparison of factors which condition choice of a hibernation place may have significant meaning in practice of protection of bats and their habitats. There is a possibility of simulation of microclimate of an underground system and consequently a refugioclimate to the needs of hibernation of a given bat species What is new The fact that choice of hibernation of both a bat species and a strategy is dependant on physical parameters of refugioclimate was not taken into account before. So far there was no simultaneous analysis of all (3) parameters (T s ; Rh; v) of refugioclimate and substrate on which bats winter (λ degree of thermal conductivity). 113

114 The work adopted proper methodology of measurement of refugioclimate It has been proved that the choice of hibernation place of a bat species and its strategy is dependant on physical conditions of refugioclimate such as T s ; Rh; v and thermal conductivity of substrate (λ) on which it hibernates. It has been demonstrated that each of the examined species possesses its own scope of physical quantities of a refuge which is optimal during hibernation, which quantities supplement one another and which distinguish one species from another. Wet Kata cooling power degrees were used in order to achieve single values of the analysed parameters such as temperature and relative air humidity as well as air flow velocity. For the first time empirical equations of three variables have been made which determine choice of hibernation places and strategies for analysed species. 114

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129 Wojtaszyn G., Rutkowski T., Stephan W., Wiewióra D., Jaros R Największe w Polsce zimowisko mopka Barbastella barbastellus (Schreber, 1774). (OKC Pokrzywna. 4 6 listopad). materiały konferencyjne.: 22. Wołoszyn B. 1976: Wyniki badań nad termiką i wilgotnością powietrza w Jaskiniach Gór Świętokrzyskich. Rocznik Swietokrzyski, Prace Geograficzne t. V. Wołoszyn B., W Zimowe spisy nietoperzy w Polsce Wyniki i ocena skuteczności. Publikacje Centrum Informacji Chiropterologicznej ISEZ PAN Kraków.: Wołoszyn B.W Nietoperze Polski. Występowanie, środowisko, status ochronny. Publikacje Centrum Informacji Chiropterologicznej ISEZ PAN, Kraków. Wołoszyn B. W Ekologiczne aspekty ochrony hibernacji nietoperzy (jaskinie i jaskinio podobne schroniska) - w europejskiej perspektywie. In. red Wołoszyn B.W., Yagt-Yazykova E., Kuśnierz A. Wpływ środowiskowych warunków na wybór hibernaculum przez nietoperze.: Yoshino M. 1975: Climate in a small area. University Tokyo Press, Tokyo. Zelinka J. 2002: Microclimatic reserch the Slovakian shov caves. Acta Carsologica. Vol. 31 No. 1. Ljubljana.: Zima J., Kovařík M., Gaisler J., Řehák Z., Zukal J Dynamics of the number of bats hibernating in the Moravian karst in 1983 to Folia Zool. 43.: Zukal, J., Berková H., Řehák Z Activity and shelter selection by Myotis myotis and Rhinolophus hipposideros hibernating in the Kateřinská cave (Czech Republic). Mammalian Biology. 70.:

130 APPENDIX NO. 1 Devices used during research Several measuring instruments were used for research in the current publication: Gas parameter gauge which is designed for measuring humidity (0 100 %), temperature ( C) and absolute pressure of gases ( hpa). The gauge is powered by built-in accumulators. Fully charged accumulators last for 10 hours of work of the device. In the gauge a high quality measuring probe was applied, which allowed to perform measurements from a certain distance. Heat and humidity sensors placed at the end of the probe are protected from damage and dust by exchangeable shield. A piezoresistive sensor working in a bridge system was used for measuring pressure. The sensor was placed inside the device. Pressure signal is supplied to the gauge through a stub pipe placed in a side wall of the casing. Each gauge is calibrated and thermally compensated within the scope C. One should remember about periodic control. Technical data ranges of measurement temperature, C relative humidity, % barometric pressure, hpa resolving power of indications temperature, C 0.1 relative humidity, % 0.1 barometric pressure, hpa 1 increased unreliability of indications, 95% temperature, C ± 0.5 relative humidity, % ± 1.5 barometric pressure, hpa ± 1 130

131 Thermo-anemometer a gauge for measuring velocity and temperature Thermo-anemometer is a portable digital gauge used to determine velocity and temperature of air. It is used both in air-conditioning and ventilation for determining air velocity. The measuring system of the probe is comprised of: digitally controlled constant temperature anemometric (CTA) bridge, precise 18-bit analog-to-digital converter and a microprocessor which controls operation of the system. The gauge possesses a precise system of thermal compensation of the bridge which provides independence of indications of velocity in a broad range of changes of temperature in flow. Impulse converters were applied in the system which reduced to minimum energy consumption of the system and completely eliminated the phenomenon of overheating of electronic elements. Technical data ranges of measurement velocity, m/s 0,01 20 temperature, C resolving power of indications velocity, m/s 0,01 temperature, C 0,1 increased unreliability of indications, 95% velocity, m/s ± (0,01 + 4% of measured value) temperature, C ± 0. Vane anemometer Measurements were performed in determined points with the use of vane anemometer AR-2. The measuring range was 0,02 ms -1 (+/- 0,01-0,02 ms -1 ), which allowed to know general ( background ) conditions prevailing inside the underground system. The measuring time was 1 minute and averaged result was read on the scale of the device. 131

132 Fig. 1. Measuring probe which allows to perform measurements from a certain distance (SENSOTRON). Table 1. Dry-bulb temperature of refugioclimate of Barbastella barbastellus depending on the month. Month Mean Standard deviation Minimum Lower quartile Median Upper quartile Maximum Significance Dry-bulb temperature T s December 7,58 2,89 1,30 5,70 8,50 9,60 12,40 January 8,37 0,73 7,00 7,80 8,45 8,80 9,70 <0,0001 February 9,08 1,51 5,50 8,80 9,50 10,05 11,10 132

133 Table 2. Detailed multiple comparison between months of hibernation of Barbastella barbastellus Dry-bulb temperature T s December January February December 1,0000 0,0009 January 1,0000 <0,0001 February 0,0009 <0,0001 Table 3. Eigenvalues of physical factors T s, Rh and v of hibernation of Barbastella barbastellus No, of value Eigenvalue % of total variance Cumulative eigenvalues Cumulative % 1 1,18 39,48 1,18 39,48 2 0,95 31,72 2,14 71,20 3 0,86 28,80 3,00 100,00 133

134 Table 5. Diversification of chosen bat species in relation to: T s ; Rh and v. Species parameter RHH MYD MYN MYM BAR PAR n=558 n=2531 n=444 n=1548 n=220 n=537 Average T, 0 C Basic statistics of dry-bulb temperature of air, o C 8,21 9,08 9,58 9,11 8,60 7,01 min T 0 C 5,3 2 5,9 3,1 1,3 0,7 max T 0 C 9,7 11,9 11,9 11,9 12,4 11,5 s.d. 1,13 1,35 1,50 1,50 1,60 2,35 Standard error 0,05 0,03 0,07 0,04 0,11 0,10 Trust -95% 8,11 9,03 9,44 9,04 8,38 6,81 Trust +95% 8,31 9,13 9,72 9,20 8,81 7,21 Lower quartile 7,00 8,20 8,55 8,20 7,90 5,70 Median 8,2 9,1 9,9 9,5 8,8 7,1 Upper quartile Significance Average Rh, % 8,80 10,00 10,40 10,20 9,60 8,80 a b d b a C < 0,0001 Basic statistics of relative humidity of air, Rh% 99,08 87,65 90,08 89,59 77,04 87,15 min Rh, % 98 53,6 58,4 56,8 56,4 56,7 max Rh, % ,1 99, ,8 100 s.d. 0,90 6,38 4,84 5,48 6,58 8,94 Standard error 0,04 0,13 0,23 0,12 0,44 0,39 134

135 Trust -95% Trust +95% Lower quartile 99,01 87,41 89,63 89,31 76,16 86,40 99,12 87,91 90,53 89,86 77,91 87,91 98,00 83,10 87,60 86,80 72,00 80,00 Median 98,5 88,8 90,3 90,3 77,0 86,3 Upper quartile Significance Average V, m/s 100,00 92,90 93,00 93,00 81,10 97,00 d a b b c A < 0,0001 Basic statistics of air flow velocity v, m/s 0,06 0,26 0,14 0,25 0,24 0,20 min V 0,03 0,01 0 0,01 0,01 0,01 max V 0,1 1,42 0,9 1,42 1,25 1,42 s,d, 0,03 0,35 0,21 0,33 0,28 0,25 Standard error Trust -95% Trust +95% Lower quartile 0,001 0,007 0,010 0,008 0,018 0,011 0,06 0,25 0,12 0,23 0,20 0,18 0,06 0,27 0,16 0,26 0,27 0,23 0,04 0,04 0,02 0,03 0,05 0,07 Median 0,08 0,07 0,04 0,07 0,12 0,08 Upper quartile Significance 0,09 0,33 0,13 0,30 0,37 0,28 c b d ab ab A < 0,

136 Table 6. Detailed multiple comparison between chosen species Dry-bulb temperature T s BAR MYD MYM MYN PAR RHH BAR 0,0097 0,0041 <0,0001 <0,0001 0,0798 MYD 0,0097 0,9875 <0,0001 <0,0001 <0,0001 MYM 0,0041 0,9875 0,0001 <0,0001 <0,0001 MYN <0,0001 <0,0001 0,0001 <0,0001 <0,0001 PAR <0,0001 <0,0001 <0,0001 <0,0001 <0,0001 RHH 0,0798 <0,0001 <0,0001 <0,0001 <0,0001 Relative humidity BAR MYD MYM MYN PAR RHH BAR <0,0001 <0,0001 <0,0001 <0,0001 <0,0001 MYD <0,0001 <0,0001 <0,0001 0,8157 <0,0001 MYM <0,0001 <0,0001 0,8316 <0,0001 <0,0001 MYN <0,0001 <0,0001 0,8316 <0,0001 <0,0001 PAR <0,0001 0,8157 0, , <0,0001 RHH <0,0001 <0,0001 <0,0001 <0,0001 <0,0001 Air flow BAR MYD MYM MYN PAR RHH BAR 0,9596 0,9985 0,0091 0,9007 <0,

137 Dry-bulb temperature T s BAR MYD MYM MYN PAR RHH BAR 0,0097 0,0041 <0,0001 <0,0001 0,0798 MYD 0,0097 0,9875 <0,0001 <0,0001 <0,0001 MYM 0,0041 0,9875 0,0001 <0,0001 <0,0001 MYN <0,0001 <0,0001 0,0001 <0,0001 <0,0001 MYD 0,9596 0,8644 <0,0001 0,0385 <0,0001 MYM 0,9985 0,8644 <0,0001 0,2052 <0,0001 MYN <0,0001 <0,0001 <0,0001 0,0115 0,0045 PAR 0,9007 0,0385 0,2052 0,0115 <0,0001 RHH <0,0001 <0,0001 <0,0001 0,0045 <0,0001 Table 7. Choice of New Kata cooling power degrees by bats in hibernation places. Mean Standard deviation Minimum Lower quartile Median Upper quartile Maximum Significance Kata cooling power degrees BAR 1021,71 253,92 660,54 823,02 950, , ,38 ac MYD 972,69 277,47 575,46 771,50 890, , ,59 a MYM 957,37 260,71 575,46 755,73 891, , ,59 a MYN 833,14 226,04 439,53 635,91 777,22 961, ,04 b < 0,0001 PAR 1033,38 202,05 620,24 911,80 966, , ,59 c RHH 823,15 84,84 543,90 747,99 848,33 917,89 969,38 b 137

138 Table 8. Detailed Tukey s multiple comparison test between examined species Kata cooling power degrees BAR MYD MYM MYN PAR RHH BAR 0,3076 0,0740 <0,0001 0,9965 <0,0001 MYD 0,3076 0,5253 <0,0001 0,0010 <0,0001 MYM 0,0740 0,5253 <0,0001 <0,0001 <0,0001 MYN <0,0001 <0,0001 <0,0001 <0,0001 0,9913 PAR 0,9965 0,0010 <0,0001 <0,0001 <0,0001 RHH <0,0001 <0,0001 <0,0001 0,9913 <0,0001 Table 9. Range of choice of wet Kata cooling power degrees of lesser horseshoe bat Rhinolophus hipposideros Descriptive statistics Mean Standard deviation Minimum Lower quartile Median Upper quartile Maximum Kata cooling power degrees 19,59 2,02 12,95 17,55 20,20 21,85 23,08 138

139 Table 10. Diversification of chosen strategies for BAR: T s ; Rh; V. Strategy Ia Ib Ib Ic IIa IIb IIc parameter concrete rubber brick concrete n = 6 n = 150 n = 36 n = 8 n= n = 16 n = 4 Basic statistics of dry-bulb temperature of air, o C Average T, 0 C 8,73 8,93 7,61 10,67 7,51 5,03 min T 0 C 7,2 6,00 3,5 9,3 6,0 1,3 max T 0 C 9,6 11,1 9,7 12,4 9,5 8,6 s.d. 0,89 1,20 1,83 1,13 0,98 3,71 Standard error 0,32 0,10 0,30 0,40 0,24 1,85 Trust -95% 7,80 8,73 6,99 9,73 6,99-0,87 Trust +95% 9,66 9,12 8,23 11,61 8,03 10,92 Lower quartile 8,50 8,30 6,40 10,05 7,00 1,85 Median 8,8 9,2 8,1 10,4 7,7 5,1 Upper quartile 9,50 9,90 8,85 11,40 8,00 8,20 Significance <0,0001 Basic statistics of relative humidity of air, Rh% Average Rh, % 71,52 77,20 73,42 85,52 80,4 81,4 min Rh, % 66,4 56,4 67,5 79,1 69,8 74,5 max Rh, % 74,5 91,8 80,5 90,1 90,8 90,1 s.d. 3,12 6,83 3,60 4,13 4,35 6,90 Standard error Trust -95% 1,27 0,56 0,60 1,46 1,09 3,45 68,24 76,10 72,20 82,07 78,08 70,42 139

140 Trust +95% Lower quartile 74,78 78,30 74,64 88,98 82,72 92,38 69,60 72,80 70,50 82,85 78,60 76,00 Median 72,1 78,0 73,2 86,6 79,7 80,5 Upper quartile 74,50 81,10 76,50 88,05 82,25 86,80 Significance <0,0001 Basic statistics of air flow velocity v, m/s Average V, m/s 0,57 0,15 0,42 0,73 0,17 0,51 min V 0,50 0,01 0,02 0,33 0,02 0,35 max V 0,67 0,92 1,25 1,17 0,92 0,63 s.d. 0,08 0,19 0,32 0,40 0,26 0,14 Standard error 0,03 0,02 0,05 0,14 0,06 0,72 Trust -95% 0,49 0,12 0,31 0,40 0,04 0,28 Trust +95% 0,66 0,18 0,53 1,06 0,31 0,74 Lower quartile 0,50 0,05 0,13 0,37 0,02 0,39 Median 0,56 0,11 0,41 0,63 0,03 0,53 Upper quartile 0,67 0,13 0,63 1,17 0,30 0,63 Significance <0,0001 Correlation coefficient T; Rh; V r T, Rh 0, ,5373-0,360,1-0,9955-0,4799-0,9561 r T, V -0,8094 0, ,0573-0,3141-0,0145-0,9628 r Rh, V -0,1233-0,6335 0, ,3184 0, ,

141 Table 11. Detailed multiple comparison between strategies of Barbastella barbastellus Dry-bulb temperature Ia Ib - concrete Ib - rubber Ic IIb IIc Ia 1,0000 1,0000 0,1657 1,0000 1,0000 Ib - concrete 1,0000 0,0010 0,0242 0,0004 0,1192 Ib - rubber 1,0000 0,0010 <0,0001 1,0000 1,0000 Ic 0,1657 0,0242 <0,0001 <0,0001 0,0007 IIb 1,0000 0,0004 1,0000 <0,0001 1,0000 IIc 1,0000 0,1192 1,0000 0,0007 1,0000 Relative humidity Ia Ib - concrete Ib - rubber Ic IIb IIc Ia 0,2073 1,0000 0,0007 0,0176 0,2780 Ib - concrete 0,2073 0,0082 0,0194 0,6677 1,0000 Ib - rubber 1,0000 0,0082 0,0001 0,0015 0,4656 Ic 0,0007 0,0194 0,0001 1, ,0000 IIb 0,0176 0,6677 0,0015 1,0000 1,0000 IIc 0,2780 1,0000 0,4656 1,0000 1,0000 Air flow Ia Ib - concrete Ib - rubber Ic IIb IIc Ia 0,0038 1,0000 1,0000 0,0041 1,0000 Ib - concrete 0,0038 0,0001 0,0005 1,0000 0,

142 Dry-bulb temperature Ia Ib - concrete Ib - rubber Ic IIb IIc Ia 1,0000 1,0000 0,1657 1,0000 1,0000 Ib - concrete 1,0000 0,0010 0,0242 0,0004 0,1192 Ib - rubber 1,0000 0,0010 <0,0001 1,0000 1,0000 Ic 0,1657 0,0242 <0,0001 <0,0001 0,0007 Ib - rubber 1,0000 0,0001 1,0000 0,0081 1,0000 Ic 1,0000 0,0005 1,0000 0,0010 1,0000 IIb 0,0041 1,0000 0,0081 0,0010 0,0610 IIc 1,0000 0,0928 1,0000 1,0000 0,0610 Table 12. Choice of wet Kata cooling power degrees depending on strategy of Barbastella barbastellus Mean Standard deviation Minimum Lower quartile Median Upper quartile Maximum Significance Wet Kata cooling power degrees Ia 32,37 1,62 31,05 31,35 31,78 32,81 35,44 Ib - concrete 22,46 4,81 15,73 19,04 22,15 23,34 37,30 Ib - rubber 29,10 6,35 17,54 24,13 31,60 33,81 37,84 Ic 30,17 4,25 25,54 26,13 29,85 34,63 34,63 IIb 22,73 6,25 17,60 18,06 19,96 28,04 37,30 IIc 33,98 1,00 32,90 33,17 33,94 34,78 35,12 < 0,

143 Table 13. Detailed multiple comparison between strategies of choice of wet Kata cooling power degrees by Barbastella barbastellus Wet Kata cooling power degrees Ia Ib - concrete Ib - rubber Ic IIb IIc Ia 0,0077 1,0000 1,0000 0,0153 1,0000 Ib - concrete 0,0077 <0,0001 0,0058 1,0000 0,0150 Ib - rubber 1,0000 <0,0001 1,0000 0,0078 1,0000 Ic 1,0000 0,0058 1,0000 0,0163 1,0000 IIb 0,0153 1,0000 0,0078 0,0163 0,0200 IIc 1,0000 0,0150 1,0000 1,0000 0,0200 Table 14. Diversification of chosen strategies for MYD: T s ; Rh; V. Strategy Ia Ib Ic IIa IIb IIc parameter n=1 n=2058 n=3 n=469 Basic statistics of dry-bulb temperature of air, o C Average T, o C 9,6 9,13 7,83 8,86 min T 2,0 6,8 2,1 max T 11,9 9,9 11,9 s.d. 1,34 1,79 1,36 Standard error 0,03 1,03 0,06 143

144 Trust -95% 9,07 3,39 8,74 Trust +95% 9,19 12,28 8,98 Lower quartile 8,20 6,80 7,90 Median 9,2 6,8 8,8 Upper quartile 10,10 9,90 10,00 Significance 0,0029 Basic statistics of relative humidity of air, Rh% Average Rh, % 94,7 87,58 94,00 87,93 min Rh 53, ,2 max Rh 99, ,1 s.d. 6,44 5,29 6,09 Standard error Trust -95% Trust +95% Lower quartile 0,14 3,06 0,28 87,31 80,86 87,38 87,87 107,15 88,48 83,10 88,00 88,00 Median 88, ,8 Upper quartile 92,90 83,10 93,00 Significance 0,1228 Basic statistics of air flow velocity v, m/s Average V, m/s 0,02 0,26 0,29 0,25 144

145 min V 0,01 0,06 0,01 max V 1,42 0,75 1,42 s.d. 0,36 0,40 0,34 Standard error Trust -95% Trust +95% Lower quartile 0,01 0,23 0,02 0,25-0,70 0,22 0,28 1,28 0,28 0,04 0,06 0,04 Median 0,07 0,06 0,07 Upper quartile 0,33 0,75 0,33 Significance 0,6425 Correlation coefficient T; Rh; V r T, Rh 0, ,9820 0,002,63 r T, V -0,2603 1,0-0,2308 r Rh, V 0, ,9820 0,

146 Table 15. Detailed multiple comparison between strategies of Myotis daubentonii Dry-bulb temperature Ia Ib Ic IIb Ia 1,0000 1,0000 1,0000 Ib 1,0000 0,8182 0,0037 Ic 1,0000 0,8182 1,0000 IIb 1,0000 0,0037 1,0000 Relative humidity Ia Ib Ic IIb Ia 0,7953 1,0000 0,8575 Ib 0,7953 0,4942 1,0000 Ic 1,0000 0,4942 0,5726 IIb 0,8575 1,0000 0,5726 Air flow Ia Ib Ic IIb Ia 1,0000 1,0000 1,0000 Ib 1,0000 1,0000 1,0000 Ic 1,0000 1,0000 1,0000 IIb 1,0000 1,0000 1,

147 Table 16. Choice of wet Kata cooling power degrees depending on strategy of Myotis daubentonii Mean Standard deviation Minimum Lower quartile Median Upper quartile Maximum Significance Kata cooling power degrees Ia 16,14 16,14 16,14 16,14 16,14 16,14 Ib 23,13 6,64 13,70 18,37 21,20 27,00 40,73 Ic 24,35 6,22 20,69 20,69 20,82 31,54 31,54 IIb 23,28 6,48 13,70 18,37 21,20 27,09 40,73 0,5523 Table 17. Detailed multiple comparison between choice of strategy of Myotis daubentonii and Kata cooling power degrees were performed with the use of Dunn s test Kata cooling power degrees Ia Ib Ic IIb Ia 1, , , Ib 1, , , Ic 1, , , IIb 1, , ,

148 Table 18. Diversification of chosen strategies for MYN: T s ; Rh; V. Strategy Ia Ib Ic IIa IIb IIc parameter concrete concrete n=273 n=171 Basic statistics of dry-bulb temperature of air, o C Average T, o C 9,80 9,22 min T 5,9 6,0 max T 11,9 11,9 s.d. 1,45 1,51 Standard error Trust -95% 0,09 0,12 9,63 8,99 Trust +95% 9,98 9,45 Lower quartile 9,10 7,90 Median 10,1 9,7 Upper quartile 10,70 10,30 Significance <0,0001 Basic statistics of relative humidity of air, Rh% Average Rh, % 90,49 89,42 min Rh 58,4 72,7 max Rh 99,1 99,1 s.d. 4,48 5,30 148

149 Standard error Trust -95% 0,27 0,41 89,96 88,62 Trust +95% Lower quartile 91,02 90,22 88,80 86,80 Median 90,9 89,9 Upper quartile 93,00 92,90 Significance 0,0238 Basic statistics of air flow velocity v, m/s Average V, m/s 0,15 0,12 min V 0,01 0,1 max V 0,9 0,9 s.d. 0,22 0,20 Standard error Trust -95% 0,01 0,02 0,12 0,09 Trust +95% Lower quartile 0,17 0,15 0,01 0,02 Median 0,04 0,04 149

150 Upper quartile 0,18 0,09 Significance 0,2467 Correlation coefficient T; Rh; V r T, Rh r T, V r Rh, V Table 19. Choice of wet Kata cooling power degrees depending on strategy of Myotis nattereri Mean Standard deviation Minimum Lower quartile Median Upper quartile Maximum Significance Kata cooling power degrees Ib IIb

151 Table 20. Diversification of chosen strategies for MYM: T s ; Rh; V. Strategy Ia Ib Ib IIa IIb IIb IIb IIb parameter concrete dolomite concrete 2-20 concrete concrete concrete ^80 n=4 n=519 n=42 n=10 n=579 n=169 n=147 n=68 Basic statistics of dry-bulb temperature of air, o C Average T, o C 9,2 9,36 5,32 8,69 9,16 9,30 9,43 8,97 min T 7,8 6,0 3,1 6,0 6,0 6,0 6,0 6,0 max T 10,7 11,9 8,1 10,6 11,9 11,7 11,3 10,7 s.d. 1,24 1,20 1,73 1,44 1,20 1,26 1,23 1,13 Standard error 0,62 0,05 0,27 0,46 0,05 0,10 0,10 0,14 Trust -95% 7,23 9,26 4,78 7,66 9,06 9,11 9,23 8,70 Trust +95% 11,17 9,46 5,86 9,72 9,26 9,49 9,63 9,24 Lower quartile 8,25 8,30 3,40 7,90 8,20 8,20 8,30 8,55 Median 9,2 9,6 5,5 8,3 9,5 9,7 9,9 9,4 Upper quartile 10,15 10,40 6,10 9,80 10,00 10,20 10,50 9,70 Significance <0,0001 Basic statistics of relative humidity of air, Rh% Average Rh, % 78,2 88,84 97,76 88,3 88,79 90,46 91,68 89,79 min Rh 56,8 65,8 96,0 84,3 66,2 76,5 82,0 83,1 max Rh 95,0 99,1 100,0 97,8 99,1 97,8 97,8 97,8 s.d. 19,55 5,34 1,08 4,65 5,55 4,65 3,98 3,71 Standard error 9,78 0,23 0,17 1,47 0,23 0,36 0,33 0,45 151

152 Trust -95% Trust +95% Lower quartile 47,16 88,38 97,43 85,00 88,34 89,75 91,03 88,90 109,38 89,30 98,09 91,63 89,24 91,17 92,34 90,69 61,70 84,50 97,00 84,30 84,50 88,80 90,00 86,80 Median 80,6 90,0 98,0 86,8 90,0 90,0 91,8 91,1 Upper quartile 94,85 93,00 98,00 91,80 93,00 93,00 93,00 93,00 Significance <0,0001 Basic statistics of air flow velocity v, m/s Average V, m/s 0,04 0,24 0,06 0,38 0,23 0,22 0,41 0,26 min V 0,02 0,01 0,04 0,04 0,01 0,01 0,01 0,02 max V 0,08 1,42 0,09 0,75 1,42 1,37 1,37 1,22 s.d. 0,03 0,33 0,01 0,30 0,31 0,30 0,40 0,31 Standard error 0,01 0,01 0,002 0,09 0,01 0,02 0,03 0,04 Trust -95% -0,003 0,21 0,05 0,17 0,21 0,17 0,34 0,19 Trust +95% 0,09 0,27 0,06 0,60 0,26 0,26 0,47 0,34 Lower quartile 0,02 0,03 0,05 0,07 0,03 0,02 0,07 0,03 Median 0,04 0,07 0,06 0,44 0,07 0,07 0,30 0,20 Upper quartile 152 0,07 0,30 0,06 0,65 0,30 0,30 0,97 0,27 Significance <0,0001 Correlation coefficient T; Rh; V r T, Rh 0, , ,2446 0, ,0175-0,0685-0,3326-0,2168 r T, V -0,9258-0,1294 0, ,3429-0,1842-0,2263 0, ,0643 r Rh, V -0,7981 0, , ,3429 0, , , ,35969

153 Table 21. Detailed multiple comparison between choice of strategy of Myotis myotis Dry-bulb temperature Ia Ib - concrete Ib - dolomite IIa IIb IIb IIb IIb - >80 Ia 1,0000 0,1163 1,0000 1,0000 1,0000 1,0000 1,0000 Ib - concrete 1,0000 <0,0001 1,0000 0,1225 1,0000 1,0000 0,2987 Ib - dolomite 0,1163 <0,0001 0,0151 <0,0001 <0,0001 <0,0001 <0,0001 IIa 1,0000 1,0000 0,0151 1,0000 1,0000 1,0000 1,0000 IIb ,0000 0,1225 <0,0001 1,0000 0,9975 0,0067 1,0000 IIb ,0000 1,0000 <0,0001 1,0000 0,9975 1,0000 0,4911 IIb ,0000 1,0000 <0,0001 1,0000 0,0067 1,0000 0,0200 IIb - >80 1,0000 0,2987 <0,0001 1,0000 1,0000 0,4911 0,0200 Relative humidity Ia Ib - concrete Ib - IIa IIb - dolomite 2-20 IIb IIb IIb - >80 Ia 1,0000 0,0215 1,0000 1,0000 1,0000 1,0000 1,0000 Ib - concrete 1,0000 <0,0001 1,0000 1,0000 0,0974 <0,0001 1,0000 Ib - dolomite 0,0215 0,0000 <0,0001 <0,0001 <0,0001 <0,0001 <0,0001 IIa 1,0000 1,0000 <0,0001 1,0000 1,0000 0,6673 1,0000 IIb ,0000 1,0000 <0,0001 1,0000 0,2559 <0,0001 1,0000 IIb ,0000 0,0974 <0,0001 1,0000 0,2559 0,7372 1,0000 IIb ,0000 <0,0001 <0,0001 0,6673 <0,0001 0,7372 0,1802 IIb - >80 1,0000 1,0000 <0,0001 1,0000 1,0000 1,0000 0,1802 Air flow Ia Ib - concrete Ib - dolomite IIa IIb IIb IIb IIb - >80 Ia 1,0000 1,0000 0,9276 1,0000 1,0000 0,7525 1,0000 Ib - concrete 1,0000 0,2140 1,0000 1,0000 1,0000 0,0001 1,0000 Ib - dolomite 1,0000 0,2140 0,1256 0,3203 1,0000 <0,0001 0,0548 IIa 0,9276 1,0000 0,1256 1,0000 0,9730 1,0000 1,0000 IIb ,0000 1,0000 0,3203 1,0000 1,0000 <0,0001 1,0000 IIb ,0000 1,0000 1,0000 0,9730 1,0000 <0,0001 1,0000 IIb ,7525 0,0001 <0,0001 1,0000 <0,0001 <0,0001 1,0000 IIb - >80 1,0000 1,0000 0,0548 1,0000 1,0000 1,0000 1,

154 Table 22. Choice of wet Kata cooling power degrees depending on strategy of Myotis myotis Mean Standard deviation Minimum Lower quartile Median Upper quartile Maximum Significance Kata cooling power degrees Ia 18,93 3,66 15,50 15,82 18,68 22,03 22,85 Ib - concrete Ib - dolomite 22,55 6,21 13,70 17,36 20,94 26,35 40,73 21,55 1,33 19,36 20,82 21,61 22,35 24,63 IIa 26,73 7,29 18,13 18,91 29,14 32,91 35,46 IIb ,62 6,15 13,70 17,94 21,01 26,47 40,73 IIb IIb ,94 6,32 14,07 16,03 20,80 26,35 35,46 25,14 6,71 14,77 18,84 26,38 32,41 35,46 IIb - >80 23,65 5,81 16,49 17,99 24,12 26,35 38,00 0,0001 Table 23. Detailed multiple comparison of choice between strategies of Myotis myotis Kata cooling power degrees Ia Ib - concrete Ib - dolomite IIa IIb IIb IIb IIb - >80 Ia 1,0000 1,0000 1,0000 1,0000 1,0000 1,0000 1,0000 Ib - concrete Ib - dolomite 1,0000 1,0000 1,0000 1,0000 1,0000 0,0004 1,0000 1,0000 1,0000 1,0000 1,0000 1,0000 0,8618 1,

155 IIa 1,0000 1,0000 1,0000 0,8079 1,0000 1,0000 1,0000 IIb ,0000 1,0000 1,0000 0,8079 1,0000 0,0002 0,5553 IIb ,0000 1,0000 1,0000 1,0000 1,0000 0,0008 1,0000 IIb ,0000 0,0004 0,8618 1,0000 0,0002 0,0008 1,0000 IIb - >80 1,0000 1,0000 1,0000 1,0000 0,5553 1,0000 1,0000 Table 24. Diversification of chosen strategies for PAR: T s ; Rh; V. Strategy Ia Ib Ic IIa IIb IIc parameter n=12 n=519 Basic statistics of dry-bulb temperature of air, o C Average T, o C 4,78 7,03 min T 0,7 0,7 max T 11,5 11,3 s.d. 3,50 2,29 Standard error Trust -95% 1,01 0,10 2,55 6,81 Trust +95% 6,70 7,22 Lower quartile 3,20 6,00 3,90 Median 3,40 7,1 3,95 Upper quartile 5,40 8,80 4,00 Significance 0,

156 Basic statistics of relative humidity of air, Rh% Average Rh, % 96,33 86,99 min Rh 93,0 56,7 max Rh ,0 s.d. 2,46 8,90 Standard error Trust -95% Trust +95% Lower quartile 0,71 0,39 94,76 86,22 97,89 87,76 95,00 79,70 97,00 Median 96,0 85,5 97,50 Upper quartile 98,00 97,00 98,00 Significance 0,0013 Basic statistics of air flow velocity v, m/s Average V, m/s 0,11 0,21 min V 0,02 0,01 max V 0,38 1,42 s.d. 0,13 0,25 Standard error Trust -95% Trust +95% 0,37 0,01 0,03 0,18 0,19 0,23 156

157 Lower quartile 0,03 0,07 0,03 Median 0,08 0,08 0,06 Upper quartile 0,09 0,28 0,09 Significance 0,2239 Correlation coefficient T; Rh; V r T, Rh -0,2617-0,6595 r T, V 0, ,12139 r Rh, V -0,7584-0,0610 Table 25. Dry-bulb temperature, relative humidity and air flow depending on the strategy of Plecotus auritus Dry-bulb temperature Ia Ib Ic Ia 0,0116 1,0000 Ib 0,0116 0,2069 Ic 1,0000 0,2069 Relative humidity Ia Ib Ic Ia 0,0059 1,00000 Ib 0,0059 0,1582 Ic 1,0000 0,1582 Air flow Ia Ib Ic Ia 0,5098 1,

158 Dry-bulb temperature Ia Ib Ic Ia 0,0116 1,0000 Ib 0,0116 0,2069 Ic 1,0000 0,2069 Relative humidity Ib 0,5098 0,8788 Ic 1,0000 0,8788 Table 26. Choice of wet Kata cooling power degrees depending on strategy of Plecotus auritus Mean Standard deviation Minimum Lower quartile Median Upper quartile Maximum Significance Kata cooling power degrees Ia 23,14 2,99 18,35 20,90 24,28 24,99 26,68 Ib 24,65 4,85 14,77 21,71 23,00 26,90 40,73 0,8027 Ic 22,33 2,74 20,39 20,39 22,33 24,27 24,27 158

159 Table 27. Kata cooling power degrees depending on strategy of Plecotus auritus Kata cooling power degrees Ia Ib Ic Ia 1, , Ib 1, , Ic 1, , List of formulae of quadratic and linear functions for chosen six species of bats for average annual temperature of +8 0 C. strategy species RHH Ia Quadratic function for majority (if all??) registered specimens Rh = 78,3142+3,1184 T+316,644 v-0,1567 T T- 14,2066 T v-1483,271 v v Linear function in the range of lower and upper quartile of variables Rh = 97,7708-0,0456 T+27,2202 v BAR Ia BAR Ib (λ 1) BAR Ib (λ 0,16) BAR IIb MYD Ib MYD IIb (λ 1) Rh = 394, ,7881 T+19,6503 v+2,3965 T T+ 47,5522 T v-411,4992 v v Rh = 110,5609-5,2788 T-52,2461 v+0,141 T T+ 7,8808 T v+12,9543 v v Rh = 61,8758+1,4684 T+51,4689 v-0,0018 T T- 7,0409 T v+9,6291 v v Rh = 51,5677+6,846 T+68,2567 v-0,3917 T T- 9,2499 T v+0,4008 v v Rh = 118,0028-8,3647 T+6,559 v+0,523 T T+ 0,0635 T v-2,2267 v v Rh = 95,713-2,0012 T-3,3507 v+0,1161 T T+ 0,666 T v+0,2346 v v Rh = 79,4785+1,0833 T-30,1263 v Rh = 91,3035-1,7913 T+12,6431 v Rh = 78,8284-0,7532 T+0,7608 v Rh = 95,8774-2,1269 T+2,7887 v Rh = 77,7761+0,9333 x+4,9312 y Rh = 85,6986+0,172 x+2,791 y MYN Ib Rh = 153, ,7292 T-30,6339 v+0,7278 T T+ 3,8154 T v-2,6111 v v Rh = 87,3325+0,3219 T+0,5162 v 159

160 MYN IIb (λ 1) Rh = 85,5426-0,2928 T+53,9946 v+0,0638 T T- 4,1104 T v-19,7282 v v Rh = 83,2374+0,618 T+3,994 v MYM Ib Rh = 120,7662-7,6628 T-0,9409 v+0,4365 T T+ 0,681 T v-2,1618 v v Rh = 83,9975+0,43 x+3,3719 y MYM IIb (λ 1) Rh = 100,2182-2,9481 T+9,0412 v+0,1734 T T+ 0,1071 T v-5,839 v v Rh = 89,6914-0,1217 x+3,9555 y PAR Ia Rh = 94,8675+0,7873 T-11,6063 v+0,0657 T T- 7,9423 T v+135,1346 v v Rh = 96,2589+0,7776 x-32,1041 y PAR Ib (λ 1) Rh = 105,4839-1,7793 T-41,8804 v-0,1091 T T+ 5,5733 T v+1,3347 v v Rh = 104,9037-2,5695 x+0,6854 y 160

161 List of acronyms and symbols used in the work Table 1. List of bats. acronyms BAR MYD MYM MYN PAR RHH Latin name Barbastella barbastellus Myotis daubentonii Myotis myotis Myotis nattereri Plecotus auritus Rhinolophus hipposideros Table 2. Physical quantities. Quantity symbol/s Quantity name/s λ Thermal conduction of materials - W m -1 K -1 p atmospheric pressure - hpa v air flow velocity m s -1 Rh relative humidity of air - % T temperature - 0 C = T s =T a T b body temperature T s T a T m T ch K a K w dry-bulb temperature 0 C= T = T a ambient temperature wet-bulb temperature 0 C temperature of hibernation surface - 0 C cooling power rate (Kata degree) wet Kata cooling power degree 161

162 Table 3. Wintering strategies symbol/s Ia Ib Ic IIa IIb IIc Description wintering strategy of a bat which winters individually (hanging freely), in which convection is the decisive factor which determines cooling wintering strategy of a bat which winters individually (hanging on a wall), in which convection and conduction are the decisive factors which determine cooling wintering strategy of a bat which winters individually (in a crevice), in which the decisive factor which determines cooling is above else conduction and to a minimum degree it is convection wintering strategy of a bat which winters in groups (hanging freely), in which convection is the decisive factor which determines cooling wintering strategy of a bat which winters in groups (hanging on a wall), in which convection and conduction are the decisive factors which determine cooling wintering strategy of a bat which winters in groups (in a crevice), in which the decisive factor which determines cooling is above else conduction and to a minimum degree it is convection 162

163 Glossary Abiotic devoid of life; non-living. Aggregation process of combining (single specimens) into a bigger whole. The phenomenon belongs to important physiological and ecological behaviours among others for hibernation of organisms. Air flow velocity the amount of air flowing through distinguished space, area or cross section in a unit of time. Anemometer a device used to measure velocity and sometimes direction of the wind. Barometer a device used to measure atmospheric pressure. Brown fatty (adipose) tissue a kind of adipose tissue in which multilocular fat cells (adipocytes) with plenty mitochondria dominate; these cells take part in processes of thermoregulation by producing heat. Cluster grouping of bats which cuddle up to one another or lie on one another. Condensation of water vapour the process of condensation of water vapour contained in the air Convection the process of heat transfer connected with macroscopic movement of matter in gas, liquid or plasma in e.g. the air. Degree of cooling - The quantity of cooling power of the atmosphere, which is intensity of cooling K, is expressed by loss of heat from 1 cm 2 of surface in 1 second. The unit of cooling intensity is Kata cooling power degree. Ecological preferendum the zone which is characterized by occurrence of a value of an environmental factor or factors optimal for a given species. Endothermy a mechanism of maintaining specific temperature of animal body, usually higher than ambient, thanks to internal sources of heat, which facilitates thermoregulation and homoiothermy. Enthalpy thermodynamic quantity equal to the sum of internal energy of air and a product of its volume and pressure. Estivation passing the summer or dry season in a dormant or torpid state. A physiological state in which pace of life processes is reduced; it occurs both in cold and warm-blooded animals; it facilitates economical management of water and energy. 163

164 Euthermia thermal state connected with change of set point of an organism. It is beneficial and favourable for the organism. The state of euthermia is fever (thanks to it immunological response develops) and anapyrexia (because it reduces loss of oxygen and limits metabolism during oxygen deficiency). Excavation empty space in a rock mass made by removing rocks during mining works. Exchange of heat transferring heat in a given medium (e.g. through the air) or between various media. Heterothermy - The mechanism of regulation of body temperature of warm-blooded animals: birds and mammals, which are capable of seasonal change of constant body temperature during torpor or hibernation. Such animals are called heterotherms. Hibernation physiological state of long-term torpor of an organism which is manifested through seasonal inhibition of life processes in some warm-blooded animals and which allows them to survive hard conditions of winter. Winter sleep may be a continuous or intermittent state. In the narrow sense of the word hibernation refers to warm-blooded animals in which when cold season starts there is a halt in activities connected with movement, significant reduction of pace of metabolism and a whole range of life functions (reduction of frequency of breathing, heart rhythm, decrease in heat production) and decrease of body temperature to the ambient temperature (however, it is often specific and constant for the hibernating animal). Hibernaculum the domicile in which an animal hibernates or overwinters; winter quarters. Homoiotherms warm-blooded animals (birds and mammals) of constant body temperature, often higher than ambient temperature. Homoiothermy ability to regulate own body temperature independent of ambient temperature fluctuations. State or ability of an organism to maintain body temperature on a constant level or fluctuate in a narrow range; it may be based on ectothermy and is connected with life in an environment of a constant, unchangeable temperature (e.g. environment of the bottom of oceans of temperature ca. 2 ); usually it is achieved by the means of endothermy and then it may be independent from significant changes of ambient temperature, the state results then from balance between producing heat (in metabolic reactions which release heat, e.g. during contraction of a muscle or burning fat in brown adipose tissue) and its loss (radiation, conduction, vaporization of water from respiratory tract and through skin, excretion). Humidity content of water (water vapour) in a solid or gaseous substance; more broadly: content of any liquid in a substance. absolute humidity the actual amount of water vapour present in a unit mass of air, expressed in grams (or kilograms) per cubic metre. air humidity content of water vapour in the air (atmosphere). 164

165 relative humidity ratio of partial pressure of water vapour contained in the air to saturation pressure, which defines maximum partial pressure of water vapour in a given temperature. Expressed in values per cent. Hypothermia the condition of a homoiothermic animal which has a lower than normal temperature Kata cooling power degree unit of amount of heat which is taken from surface area of one square centimetre in one second at a determined temperature; it is used to determine intensity of cooling. Katathermometer a device to measure intensity of cooling influence of environment, induced by common influence of temperature, velocity and humidity of the air. Measurement error variance between reading of value of a given meteorological element and real value of this element. Metabolism the totality of the synthtetic and degradative biochemical processes of living organisms. Microclimate climate of a small area of surface ranging from several up to several hundred square metres, with characteristics which differ given area from surrounding environment, e.g. climate of a field, slope, gorge, edge of wood, lakeside, crown of a tree or climate of the layer of air close to the ground (up to 2 m) over a specific homogeneous surface. interior microclimate = microclimate of an underground system = microclimate of a cave microclimatic conditions of closed spaces, inside houses, warehouses, cargo holds etc.; the term includes also natural places like e.g. caves, animal burrows etc. Mollier diagram a diagram which presents mutual relations between air temperature, humidity and enthalpy. Poikilothermy changeability of body temperature of animals (called cold-blooded animals) along with fluctuations of ambient temperature. Lack of ability to keep heat and regulate body temperature and activeness of metabolism in relation to fluctuations of temperature in the environment of the animal. Pressure physical value defined by a ratio of a force acting vertically onto a given surface to the surface area. atmospheric pressure pressure induced by air on all objects located within it; it is measured by barometers. pressure (compressibility) of saturated water vapour; water vapour pressure (WVP) partial pressure of water vapour, which at that point in time is present in the air, at which in a given temperature gas is in balance with liquid. Balance between vaporization and condensation occurs. It is expressed in hpa, Psychrometer in meteorology the most popular device for measuring air humidity. 165

166 Refugioclimate a set of physical factors which have an effect directly in a hibernation place on a bat (or other organism) (usually several cm around). Roof of an underground system - upper surface of underground excavation. Side wall side wall of an underground system (an excavation in a mine, a cave). Temperature basic quantity which determines thermodynamic state of the system; value of temperature is given in degrees depending on the adopted scale; the unit of temperature in absolute scale is kelvin (K); in meteorology popular is also Celsius scale ( C). dry-bulb temperature air temperature; temperature which is shown by a regular thermometer (i.e. dry one) wet-bulb temperature temperature which is shown by a moistened thermometer or one covered with ice. Thermal comfort it is such a state of satisfaction of a specimen (group) in thermal conditions of environment in which it feels neither warmth nor coolness. The necessary condition to feel thermal comfort is achieving the state of thermal balance of organism. Such a state is characterized by levelling the amount of heat of metabolism with energy exchanged between the organism (of a hibernating bat) and the environment. Thermal conduction of materials determines capability of a substance to conduct heat. In same conditions more heat will flow through a substance of higher thermal conduction coefficient. The substances which best conduct heat are metals, while gases are the poorest conductors. Thermal radiation electromagnetic radiation sent by any body at temperature higher than absolute zero. Thermogenesis processes connected with heat production (endothermy) necessary to maintain specific body temperature, higher than ambient temperature; in animals it is connected with movements of muscles (shivering thermogenesis) or heat production processes in brown fatty tissue (non-shivering thermogenesis). Thermoregulation thermal homeostasis, ability of organisms to maintain relatively stable body temperature, which is optimal for the course of life processes, as well as mechanisms which lead to it. It is achieved by the use of external (ectothermy) or internal (endothermy, thermogenesis) sources of heat, as well as mechanisms and structures which facilitate keeping heat and reduce its loss and if overheated they allow to lose the excess heat. social thermoregulation cuddling up to one another, which reduces heat loss. Torpor short-term state of periodic controlled decrease in body temperature and significant reduction or halt of metabolic and physiological processes as well as movement activities in result of occurrence of unfavourable environmental 166

167 conditions. Underground corridor a corridor in an underground excavation which is constructed horizontally or almost horizontally. Vaporization; evaporation process of change of state of aggregation from liquid into gaseous. 167

168