DOI: 10.5281/zenodo.7226367
title: Data from: Macro-evolutionary trade-offs as the basis for the distribution of European bats

description: We have compiled a dataset of life history traits and distribution characteristics of 30 European bat species, based on a literature study of a total of 56 primary and secondary sources. These life history traits are grouped into morphological, physiological and ecological adaptations. <em>Physiological adaptations:</em> Neonatal mass: the average weight (g) of a newborn pup, measured within five days after birth. Average litter size: the average size of a full-term litter (including stillborn pups) per female. Weaning mass: the weight (g) of a juvenile during its first flight outside the roost. Adult body mass: the average weight (g) of adult bats during the summer (between 1 May and 1 July), excluding pregnant females. Litter mass: neonatal mass * average litter size. Relative mass of neonatal to adult: neonatal mass*100 / adult weight Relative mass of litter to adult: litter mass*100 / adult weight Gestation: the length of gestation period (in days), from fertilisation to birth. When mated during autumn or winter, the sperm (or fertilised egg in <em>M. schreibersii</em>) is stored throughout the winter. On arousal from hibernation in the spring, around mid March, female bats ovulate and gestation begins. In accordance with other researchers (e.g. Altringham 1996, Entwisle <em>et al</em>. 1998), 15 March was used as the start of the gestation period, for statistical reasons we also included <em>M. schreibersii</em>. Weaning: the length of the lactation period (in days) until offspring are fully independent. After the juveniles are capable of flight, mothers continue to give their young nourishment until they are fully independent. Only when no extra nourishment is provided are the offspring considered fully weaned. Reproductive period: gestation + weaning (in days). Average age at first reproduction: The age (in days) at which 75% of the female population becomes sexual mature. Many species reproduce just before or during their first winter (at approximately 80 days old), but in some species the majority of the population postpone their sexual development. Individuals are stated to have become sexual mature if they participate in mating, have been found to be pregnant or inseminated. Observed average age: observed average age of adults in a population at a given time (in years). Longevity: the age (in years) of the oldest observed individual. The longevity can only be obtained by marking and later recapturing individuals. Most recapture data are collected in summer roosts or hibernacula. As not all species show the same fidelity to summer roost sites or can be found in hibernacula that are accessible to humans, this measure is sensitive to the chance of recapture. Minimum hibernation temperature: the minimum temperature (degrees Celsius) at which each species is observed. <em>Morphological adaptations</em> Length of forearm at birth: the length of the forearm (mm) of a newborn bat, measured between the elbow to the wrist of a folded wing. This is widely accepted as a measurement of size. Although it is not the best reflection of the length of an individual, it can be measured rapidly and accurately under field conditions. Length of forearm adult: The length of the forearm (mm) of an adult bat. Relative length of forearm of a newborn to an adult: (length of the forearm at birth*100)/ Length of forearm adult. Wing span: the length of the wings (m). The distance between the wingtips of a bat with wings extended so the leading edge is straight (including body width). Wing area: The combined area of the two wings (m<sup>2</sup>) including the entire tail membrane and the portion of the body between the wings. Wing loading: the relation between body weight, wing size and gravity (Nm<sup>-2</sup>). This measurement is related to the mean pressure on the wings. Wing loading is the weight (mass, in kg, times gravitational acceleration) divided by the wing area, i.e Wing loading = (weight adult*9.81)/ wing area. The wing load can vary significantly between geometrically similar bats. Because of such allometry, large bats have a higher wing load than smaller bats. Wing aspect ratio: the square of the wingspan divided by the wing area, i.e. Wing aspect ratio = (wingspan)<sup>2</sup> / wing area. This ratio can be interpreted as a measure of the aerodynamic efficiency of flight. A higher aspect ratio usually corresponds with greater aerodynamic efficiency (i.e. a streamlined body) and lower energy use in flight. Flight speed: The speed of flight (m/s). The speed of flight is usually measured in wind tunnel experiments or during radio-tracking. <em>Ecological adaptations </em> Maximum migration distance: the maximum observed distance (km) between the summer and winter habitat. In contrast to birds, the direction of migration in bats is not determined by the change of the seasons, but by the locations of the hibernacula. This migration distance can only be obtained by capturing, marking and later recapturing individuals. Bats often migrate across national boundaries and gathering recapture data requires international cooperation. The chance of recapture is sensitive to sample effort and local observation methods. Average migration distance: the average distance (km) between the summer and winter habitat. Most species of bats migrate both short and long distances. The same restrictions described for maximum migration distance also apply to this parameter. Echolocation type: the predominant echolocation type used by each bat species. European bats use one or sometimes a combination of the following four types of echolocations: fm-CF-fm, fm-QCF (with the QCF part dominant), FM-qcf (with the FM part dominant) and FM. For statistical reasons both FM-qcf and FM are clustered in the group FM. The FM-qcf and fm-QCF echolocations are both often loud and used to detect distant prey. FM and fm-CF-fm echolocations are softer and bats using these types of echolocation receive more detailed knowledge of their surroundings. Bats primarily use only one type of echolocation, although many can make some slight adjustments to this. Echolocation range: the maximum distance that an echolocating bat can detect a structure or object. Echolocation minimum frequency: the minimum echolocation frequency (MHz) used by each bat species. Echolocation maximum frequency: the maximum echolocation frequency (MHz) used by each bat species. Duration call (ms): the average duration (in ms) of one complete call cycle. <em>Distribution parameters</em> Northern limit of range: the most northerly observation (in latitude) of each bat species. This measurement includes anecdotal observations and observations of male bats. Northern limit of reproduction range: the most northerly observation (in latitude) of a maternity group. Note: confusion is possible between summer roosts and maternity roosts. Summer roosts are often inhabited by both males and females and less than 70% of the adult females participate in reproduction. Maternity roosts are predominantly occupied by females, and more than 70% of the adult females participate in reproduction. Southern limit of range: the most southerly observation (in latitude) of each bat species. This measurement includes anecdotal observations and observations of male bats. Southern limit of reproduction range: the most southerly observation (in latitude) of a maternity group. The same restrictions described for northern limit of reproduction range also apply to this parameter. Western limit of range: the most western observation (in longitude) of each bat species Eastern limit of range: the most eastern observation (in longitude) of each bat species Night length: The average night length (in hours) during midsummer (21<sup>st</sup> June) at the northern limit of the reproduction range. Sources: 1. Jones et al. 2009, 2. Krapp 2011, 3. Schober & Grimmberger 1997, 4. Norberg & Rayner 1987, 5. Hutterer et al. 2005, 6. Dietz et al. 2009, 7. Supplementary data from Barclay et al. 2004, 8. Wilkinson & South 2002, 9 Jones & Rydell 1994, 10. Norberg 1986, 11. Jones 1994, 12. Baagøe 1987, 13.Fleming & Eby 2003, 14. Neuweiler 2000, 15. Hayssen et al. 1993, 16. Kunz & Kurta 1987, 17. Russo & Jones 2002, 18. Brunet-Rossinni & Austad 2004, 19. Aldridge 1987, 20. Urbańczyk 1991, 21. Nagel & Nagel 1991, 22. Masing & Lutsar 2007, 23. Masing 1983, 24. Gaisler 1970, 25. Norberg 1987, 26. Baydemür & Albayrak 2006, 27. Dietz et al. 2006, 28. Sharifi 2004, 29. Kerth et al. 2001, 30. Schmidt 2005, 31. Smirnov et al. 2008, 32. Verbeek 1998, 33. Pandurkska & Beshkov 1998, 34. Harmata 1969, 35. Sachanowicz & Zub 2002, 36. Arlettaz et al. 2001, 36. Ibáñez et al. 2001, 37. Estók 2007, 38. Lohrl 1936, 39. Kunz & Hood 2000, 40. Happold & Happold 1990, 41. Rydell 1990, 42. Reiter 2004, 43. Ransome 1990, 44. Zahn 1999, 45. Deanesly & Warwick 1939, 46. Racey 1969, 47. Racey & Swift 1981, 48. Racey 1974, 49. Masing 1982, 50. Boyd & Stebbings 1989, 51. Lesiñski 1986, 52. Barak & Yom-tov 1991, 53. Arlettaz et al. 2000, 54. Gaisler et al. 1997, 55, Heise 1989, 56. Papadatou et al. 2009, 57. Unpublished data: own measurements.
description: {"references": ["Haarsma, A-J & Siepel, H. (2013). Macro-evolutionary trade-offs as the basis for the distribution of European bats. Animal Biology, 63(4): 451 \u2013 471", "Jones, K.E., Bielby, J., Cardillo, M., Fritz, S.A., O'Dell, J., Orme, C.D.L., Safi, K., Sechrest, W., Boakes, E.H., Carbone, C., Connolly, C., Cutts, M.J., Foster, J.K., Grenyer, R., Habib, M., Plaster, C.A., Price, S.A., Rigby, E.A., Rist, J., Teacher, A., Bininda-Emonds, O.R.P., Gittleman, J.L., Mace, G.M. & Purvis, A. (2009) PanTHERIA: a species-level database of life history, ecology, and geography of extant and recently extinct mammals. Ecology, 90, 2648. Ecological Archives E090-184.", "Krapp, F. (2011) Die Flederm\u00e4use Europas. Ein umfassendes Handbuch zur Biologie, Verbreitung und Bestimmung. Sonderausgabe aus dem Handbuch der S\u00e4ugetiere Europas 2011. Aula-Verlag, Wiebelsheim.", "Schober, W. & Grimmberger, E. (1997) The Bats of Europe and North America. T.F.H. Publications, Neptune City, NJ.", "Norberg, U. & Rayner, J. (1987) Ecological morphology and flight in bats: wing adaptations, flight performance, foraging strategy and echolocation. Proceedings of the Royal Society of London Series B Biological Sciences, 316, 335-427.", "Hutterer, R., Ivanova, T., Meyer-Cords, C. & Rodrigues, L. (2005) Bat Migrations in Europe: a Review of Banding Data and Literature. Federal Agency for Nature Conservation, Bonn.", "Dietz, C., Von Helversen, O. & Nill, D. (2009) Handbuch der Flederm\u00e4use Europas und Nordwestafrikas: Biologie, Kennzeichen, Gef\u00e4hrdung. Franckh-Kosmos Verlag, Stuttgart.", "Barclay, R., Ulmer, J., MacKenzie, C.J.A., Thompson, M., Olson, L., McCool, J., Cropley, E.C. & Poll, G. (2004) Variation in the reproductive rate of bats. Canadian Journal of Zoology, 82, 688- 693.", "Wilkinson, G.S. & South, J.M. (2002) Life history, ecology and longevity in bats. Aging Cell, 1, 124- 131.", "Jones, K.E. & Rydell, J. (1994) Foraging strategy and predation risk as factors influencing emergence time in echolocating bats. Philosophical Transactions, 346, 445-455.", "Baag\u00f8e, H. (1987) The Scandinavian bat fauna \u2013 adaptive wing morphology and free flight in the field. In: B.M. Fenton, P. Racey & J.M. Rayner (Eds.) Recent Advances in the Study of Bats, pp. 57-74. Cambridge University Press, Cambridge.", "Fleming, T.H. & Eby, P. (2003) Ecology of bat migration. In: T.H. Kunz & M.B. Fenton (Eds.) Bat Ecology, pp. 156-207. University of Chicago Press, Chicago, IL.", "Neuweiler, G. (2000) Reproduction and development. In: The Biology of Bats, pp. 236-261. Oxford University Press, Oxford.", "Hayssen, V., van Tienhoven, A. & van Tienhoven, A. (1993) Asdell's Patterns of Mammalian Reproduction: a Compendium of Species-Specific Data. Cornell University Press, Ithaca, NY.", "Kunz, T. & Kurta, A. (1987) Size of bats at birth and maternal investment during pregnancy. Symposia of the Zoological Society of London, 57, 79-106.", "Russo, D. & Jones, G. (2002) Identification of twenty-two bat species (Mammalia: Chiroptera) from Italy by analysis of time-expanded recordings of echolocation calls. Journal of Zoology, London, 258, 91-103.", "Brunet-Rossinni, A.K. & Austad, S.N. (2004) Ageing studies on bats: a review. Biogerontology, 5, 211-222.", "Aldridge, H.D. (1987) Turning flights in bats. Journal of Experimental Biology, 128, 419-425.", "Urbanczyk, Z. (1991) Hibernation of \u00b4 Myotis daubentonii and Barbastella barbastellus in Nietoperek bat reserve. Myotis, 29, 115-120.", "Nagel, A. & Nagel, R. (1991) How do bats choose optimal temperatures for hibernation? Comparative Biochemistry and Physiology, 99, 323-326.", "Masing, M. & Lutsar, L. (2007) Hibernation temperatures in seven species of sedentary bats (Chiroptera) in northeastern Europe. Acta Zoologica Lituanica, 17, 47-55.", "Masing, M. (1983) On the hibernation of bats in Estonia. Myotis, 20, 5-10.", "Gaisler, J. (1970) Remarks on the thermopreferendum of palearctic bats in their natural habitats. Bijdragen Dierkunde, 40, 33-35.", "Norberg, U.M. (1987) Wing form and flight mode in bats. In: M.B. Fenton, P.A. Racey & J.M.V. Rayner (Eds.) Recent Advances in the Study of Bats, pp. 43-56. Cambridge University Press, Cambridge.", "Baydem\u00fcr, N.A. & Albayrak, U. (2006) A study on the breeding biology of some bat species in Turkey (Mammalia: Chiroptera). Turkish Journal of Zoology, 30, 103-110.", "Dietz, C., Dietz, I. & Siemers, B.M. (2006) Wing measurement variations in the five European horseshoe bat species (Chiroptera: Rhinolophidae). Journal of Mammalogy, 87, 1241-1251.", "Sharifi, M. (2004) Postnatal growth in Myotis blythii (Chiroptera: Vespertilionidae). Mammalia, 68, 283-290.", "Kerth, G., Weissmann, K. & K\u00f6nig, B. (2001) Day roost selection in female Bechstein's bats (Myotis bechsteinii): a field experiment to determine the influence of roost temperature. Oecologia, 126, 1-9.", "Schmidt, C. (2005) Emergence behavior of a nursery colony of Myotis brandtii (Eversmann, 1845) in Saxony. Myotis, 43, 55-62.", "Smirnov, D.G., Vekhnik, V.P., Kurmaeva, N.M., Shepelev, A.A. & Il'in, V.Yu. (2008) Spatial structure of the community of bats (Chiroptera: Vespertilionidae) hibernating in artificial caves of Samarskaya Luka. Biological Bulletin, 35, 211-218.", "Verbeek, H. (1998) Meervleermuis Myotis dasycneme drachtig van tweeling. Lutra, 40, 89-92.", "Pandurkska, R.S. & Beshkov, A. (1998) Species diversity of bats in underground roosts of the Western Stara Planina Mts. (Bulgaria). Vespertilio, 3, 81-91.", "Harmata, W. (1969) The thermopreferendum of some species of bats. Acta Theriologica, 14, 49-62.", "Sachanowicz, K. & Zub, K. (2002) Numbers of hibernating Barbastella barbastellus and thermal conditions in military bunkers. Mammalian Biology, 67, 179-184.", "Arlettaz, R., Christe, P., Lugon, A., Perrin, N. & Vogel, P. (2001) Availability dictates the timing of parturition in insectivorous mouse-eared bats. Oikos, 95, 105-111.", "Ib\u00e1\u00f1ez, C., Juste, J., Garc\u00eda-Mudarra, J.L. & Agirre-Mendi, P.T. (2001) Bat predation on nocturnally migrating birds. Proceedings of the National Academy of Sciences of the USA, 98, 9700-9702.", "Est\u00f3k, P. (2007) Seasonal changes in the sex ratio of Nyctalus species in north-east Hungary. Acta Zoologica Academiae Scientiarum Hungaricae, 53, 89-95.", "Lohrl, H. (1936) Der winterschlaf von Nyctalus noctula \u2013 Auf grund von beobachtungen am winterschlafplatz. Zoomorphology, 32, 1-47.", "Kunz, T. & Hood, W.R. (2000) Parental care and postnatal growth in Chiroptera. In: E.G. Crichton & P.H. Krutzsch (Eds.) Reproductive Biology of Bats, pp. 415-468. Academic Press, New York, NY.", "Happold, D.C.D. & Happold, M. (1990) Reproductive strategies of bats in Africa. Journal of Zoology, 222, 557-583.", "Rydell, J. (1990) Ecology of the northern bat Eptesicus nilssoni during pregnancy and lactation. In: J. Rydell (Ed.) Ecology of the Northern Bat Eptesicus nilssoni During Pregnancy and Lactation. Department of Ecology and Animal Ecology, Lund University, Lund.", "Reiter, G. (2004) Postnatal growth and reproductive biology of Rhinolophus Hipposideros (Chiroptera: Rhinolophidae). Journal of Zoology, 262, 231-241.", "Ransome, R.D. (1990) The Natural History of Hibernating Bats. Christopher Helm Publishers, Beckenham, 235 pp.", "Zahn, A. (1999) Reproductive success, colony size and roost temperature in attic-dwelling bat Myotis myotis. Journal of Zoology (London), 247, 275-280.", "Deanesly, R. & Warwick, T. (1939) Observations on pregnancy in the common bat (Pipistrellus pipistrellus). Proceedings of the Royal Society of London, 109, 57-60.", "Racey, P.A. (1969) Diagnosis of pregnancy and experimental extension of gestation in the pipistrelle bat, Pipistrellus pipistrellus. Journal of Reproduction Fertility, 19, 465-474.", "Racey, P.A. & Swift, S. (1981) Variations in gestation length in a colony of pipistrelle bats (Pipistrellus pipistrellus) from year to year. Journal of Reproduction Fertility, 61, 123-129.", "Racey, P.A. (1974) The temperature of a pipistrelle hibernaculum. Journal of Zoology (London), 173, 260-262.", "Masing, M. (1982) On the air temperature at hibernation sites of bats. Eesti ulukid, 67, 7.", "Boyd, I.L. & Stebbings, R.E. (1989) Population changes of brown long-eared bats (Plecotus auritus) in bat boxes at Thetford Forest. Journal of Applied Ecology, 26, 101-112.", "Lesi\u00f1ski, G. (1986) Ecology of bats hibernating underground in Central Poland. Acta Theriologica, 31, 507-521.", "Barak, Y. & Yom-tov, Y. (1991) The mating system of Pipistrellus kuhli (Microchiroptera) in Israel. Mammalia, 55, 285-292.", "Arlettaz, R., Ruchet, C., Aeschimann, J., Brun, E., Genoud, M. & Vogel, P. (2000) Physiological traits affecting the distribution and wintering strategy of the bat Tadarida teniotis. Ecology, 81, 1004-1014.", "Gaisler, J., Hanak, V. & Dubngel, J. (1997) A contribution to the population ecology of Nyctalus noctula. Act. Sci. Nat., Brno, 13, 1-38.", "Heise, G. (1989) Ergebnisse reproduktionsbiologischer untersuchungen am abendsegler in der umgebung von Prezlau. Nyctalus, 3, 17-32.", "Papadatou, E., Butlin, R.K. & Pradel, R. (2009) Sex-specific roost movements and population dynamics of the vulnerable long-fingered bat, Myotis capaccinii. Biological Conservation, 142, 280-289."]}