June 2017 Version 4. A REVIEW OF EMPIRICAL DATA IN RESPECT OF EMERGENCE AND RETURN TIMES REPORTED FOR THE UK s 17 NATIVE BAT SPECIES ~~~~~~~~~~~~

Transcription

1 June 2017 Version 4 A REVIEW O EMPIRICAL DATA IN RESPECT O EMERGENCE AND RETURN TIMES REPORTED OR THE UK s 17 NATIVE BAT SPECIES ~~~~~~~~~~~~

2 CONTENTS 1. RATIONALE BARBASTELLE Barbastella barbastellus SEROTINE Eptesicus serotinus BECHSTEIN S BAT Myotis bechsteinii ALCATHOE S BAT Myotis alcathoe BRANDT S BAT Myotis brandtii DAUBENTON S BAT Myotis daubentonii WHISKERED BAT Myotis mystacinus NATTERER S BAT Myotis nattereri LEISLER S BAT Nyctalus leisleri NOCTULE Nyctalus noctula NATHUSIUS PIPISTRELLE Pipistrellus nathusii COMMON PIPISTRELLE Pipistrellus pipistrellus SOPRANO PIPISTRELLE Pipistrellus pygmaeus GREY LONG-EARED BAT Plecotus austriacus BROWN LONG-EARED BAT Plecotus auritus GREATER HORSESHOE-BAT Rhinolophus ferrumequinum LESSER HORSESHOE-BAT Rhinolophus hipposideros Report authors: Henry Andrews MSc CEcol MCIEEM & Louis Pearson BSc MSc GradCIEEM. inal proof: Leanne Butt BSc MSc ACIEEM. No part of this report may be reproduced without prior written approval of AEcol. Legal action may be taken in the event of copyright infringement. No liability is accepted in respect of the use of data, conclusions or other material contained in this report. AEcol 2016 AEcol & Andrews Ecology Ltd are registered Trademarks.

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4 REVIEW O EMERGENCE AND RETURN EMPIRICAL DATA A REVIEW O EMPIRICAL DATA IN RESPECT O EMERGENCE AND RETURN TIMES REPORTED OR THE UK s 17 NATIVE BAT SPECIES 1. RATIONALE To date, no publication or guidance document has reviewed empirical evidence in respect of emergence times, nor has any document attempted to provide the times of returns Emergence times quoted within grey literature may comprise approximations based on the author s own experiences, or cite either mean or median averages in white literature. All emergence times cited are therefore individual figures. Where the time given is an approximation the number of observations is unknown to the reader, and where they are mean averages the figure may be entirely artificial Although the mean average and standard deviation (SD) may be of scientific interest, in order to perform robust emergence and return surveillance in the context of an Ecological Impact Assessment, it is necessary to understand the temporal range across which the target species might emerge and return in that particular season. This is because it informs the duration over which each survey should be performed This review of empirical data in respect of emergence and return times reported for the UK s 17 native bat species was performed. White-papers and scientific reports were reviewed and the data collated for each species. Where appropriate, sunset and sunrise times were factored in to assess the data reported against sunset in the specific locality. Where more than one publication exists for an individual bat species, studies in the UK were preferentially selected, as were those that span both sexes and the greatest number of periods. However, in order that the reader is aware of alternative texts, other studies that were of note are included within the reference lists as see also It will be immediately apparent to the reader that many of the published accounts do not include all the salient data, there is no standardised format for the presentation of data in order that studies may be compared, and much of the data is presented in a format that confounds interpretation and how the data might be put to practical use does not appear to be considered The conclusion is that there is still room for improvement in the understanding of the temporal behaviour of all species. Immediate gains would result from the presentation of range data and SD within white literature The review was performed by Henry Andrews MSc CEcol MCIEEM and Louis Pearson BSc MSc GradCIEEM of AEcol. We would be pleased to hear from anyone who has or knows of conflicting or more detailed accounts, or has data of their own they would be prepared to share AEcol 2017

5 REVIEW O EMERGENCE AND RETURN EMPIRICAL DATA 2. BARBASTELLE Barbastella barbastellus 2.1 Emergence times Table 1 details the emergence times for the barbastelle Barbastella barbastellus reported in empirical data (Zeale et al. 2012). Table 1. Emergence times for the barbastelle Barbastella barbastellus reported in empirical data (Zeale et al. 2012). EMERGENCE TIME Mean: 24 minutes after SD: minutes (Zeale et al. 2012) after (Zeale et al. 2012) M minutes after (Zeale et al. 2012) 2.2 Return times Table 2 details the return times for the barbastelle reported in empirical data (Zeale et al. 2012). Table 2. Return times for the barbastelle Barbastella barbastellus reported in empirical data (Zeale et al. 2012). SEASON Pregnancy May/June Lactation July Weaning August RETURN TIME Mean: 194 minutes before (3 hours and 14 minutes) (Zeale et al. 2012) SD: minutes before (4 hours and 14 minutes up to 2 hours and 15 minutes) (Zeale et al. 2012) ACTUAL RANGE M All 2.3 References Zeale M, Davidson-Watts I & Jones G Home range use and habitat selection by barbastelle bats (Barbastella barbastellus): implications for conservation. Journal of Mammalogy 93(4): See also: Russo D, Cistrone L, Jones G Emergence time in forest bats: the influence of canopy closure. Acta Oecologica. 31: AEcol 2017

6 REVIEW O EMERGENCE AND RETURN EMPIRICAL DATA 3. SEROTINE Eptesicus serotinus 3.1 Emergence times Table 3 details the emergence times for the serotine Eptesicus serotinus reported in empirical data (Catto et al. 1995, Petrželkova & Zákul 2001). Table 3. Emergence times for the serotine Eptesicus serotinus reported in empirical data (Catto et al. 1995, Petrželkova & Zákul 2001). EMERGENCE TIME Mean: 11.6 minutes after (Catto et al. 1995) SD: minutes after (Catto et al. 1995) 15 minutes 45 seconds before up to 42 minutes after (Petrželkova & Zákul 2001) M 3.2 Return times Table 4 details the return times for the serotine reported in empirical data (Catto et al. 1995). Table 4. Return times for the serotine Eptesicus serotinus reported in empirical data (Catto et al. 1995). SEASON Pregnancy May/June Lactation July Weaning August RETURN TIME minutes before (5 hours and 9 minutes up to 9 minutes) (Catto et al. 1995) minutes before (3 hours and 19 minutes up to 1 hour six and minutes) (Catto et al. 1995) 66 minutes before (1 hour and six minutes) (Catto et al. 1995) M All Inferred from data presented in figures AEcol 2017

7 REVIEW O EMERGENCE AND RETURN EMPIRICAL DATA 3.3 References Catto C, Racey P & Stephenson P Activity patterns of the serotine bat (Eptesicus serotinus) at a roost in southern England. J. Zool., Lond. 235: Petrželkova K & Zákul J Emergence behaviour of the serotine bat (Eptesicus serotinus) under predation risk. Netherlands Journal of Zoology 51: See also: Jones G & Rydell J oraging strategy and predation risk as factors influencing emergence time in echolocating bats. Phil. Trans. R. Soc. Lond. B 346: AEcol 2017

8 REVIEW O EMERGENCE AND RETURN EMPIRICAL DATA 4. BECHSTEIN S BAT Myotis bechsteinii 4.1 Emergence times Table 5 details the emergence times for Bechstein s bat Myotis bechsteinii reported in empirical data (Jones & Rydell 1994, Dietz & Pir 2011). Table 5. Emergence times for Bechstein s bat Myotis bechsteinii reported in empirical data (Jones & Rydell 1994, Dietz & Pir 2011). M EMERGENCE TIME June Median: 33 minutes after (Jones & Rydell 1994) Mean: 47 minutes after (Dietz & Pir 2011) Mean: 56 minutes after (Dietz & Pir 2011) Earliest starting 30 minutes after (Jones & Rydell 1994) 2 minutes before up to 92 minutes after (2 minutes before up to 1 hour and 32 minutes after) (Dietz & Pir 2011) minutes after (25 minutes to 1 hour and 27 minutes) (Dietz & Pir 2011) 4.2 Return times Table 6 details the return times for Bechstein s bat reported in empirical data (Dietz & Pir 2011). Table 6. Return times for Bechstein s bat Myotis bechsteinii reported in empirical data (Dietz & Pir 2011). SEASON Pregnancy May/June Lactation July Weaning August RETURN TIME 60 minutes before (itzsimons et al. 2002) 60 minutes before (itzsimons et al. 2002) 60 minutes before (itzsimons et al. 2002) M All 77-9 minutes before (1 hour and 17 minutes up to 9 minutes before) (Dietz & Pir 2011) 413 minutes before up to 2 minutes after (6 hours and 53 minutes before up to 2 minutes after) (Dietz & Pir 2011) AEcol 2017

9 REVIEW O EMERGENCE AND RETURN EMPIRICAL DATA 4.3 References Dietz M & Pir J Distribution, Ecology and Habitat Selection by Bechstein s bat (Myotis bechsteinii) in Luxembourg. Ökologie der Säugetiere 6 itzsimons P, Hill D & Greenaway Patterns of habitat use by female Bechstein s bats (Myotis bechsteinii) from a maternity colony in a British woodland. School of Biological Sciences, University of Sussex Jones G & Rydell J oraging strategy and predation risk as factors influencing emergence time in echolocating bats. Phil. Trans. R. Soc. Lond. B 346: AEcol 2017

10 REVIEW O EMERGENCE AND RETURN EMPIRICAL DATA 5. ALCATHOE S BAT Myotis alcathoe 5.1 Emergence times Table 7 details the emergence times for Alcathoe s bat Myotis alcathoe reported in empirical data (observations provided by R. Baker & L. Whitfield 2016 pers comm. 16 th October, and D. Whitby 2016 pers comm. 18 th October). Table 7. Emergence times for Alcathoe s bat Myotis alcathoe reported in empirical data (observations provided by R. Baker & L. Whitfield 2016 pers comm. 16th October, and D. Whitby 2016 pers comm. 18th October). EMERGENCE TIME Mean: 1.1 minutes before (calculated from 11 observations provided by R. Baker & L. Whitfield 2016 pers comm. 16 th October, and D. Whitby 2016 pers comm. 18 th October) SD: 16.4 minutes before to 14.3 minutes after (calculated from 11 observations provided by R. Baker & L. Whitfield 2016 pers comm. 16 th October, and D. Whitby 2016 pers comm. 18 th October) 33 minutes before to 17 minutes after (calculated from 11 observations provided by R. Baker & L. Whitfield 2016 pers comm. 16 th October, and D. Whitby 2016 pers comm. 18 th October) M 5.2 Return times Table 8 details the return times for Alcathoe s bat reported in empirical data. Table 8. Return times for Alcathoe s bat Myotis alcathoe reported in empirical data. SEASON RETURN TIME ACTUAL RANGE Pregnancy May/June Lactation July Weaning August M All AEcol 2017

11 REVIEW O EMERGENCE AND RETURN EMPIRICAL DATA 6. BRANDT S BAT Myotis brandtii 6.1 Emergence times Table 9 details the emergence times for Brandt s bat Myotis brandtii reported in empirical data (Berge 2007). Table 9. Emergence times for Brandt s bat Myotis brandtii reported in empirical data (Berge 2007). EMERGENCE TIME Mean: 43.3 minutes after (Berge 2007) May/June Mean: 27.3 minutes after (Schmidt 2007) July Mean: 21.4 minutes after (Schmidt 2007) August Mean: 24.9 minutes after (Schmidt 2007) minutes after (Schmidt 2007) minutes after (Schmidt 2007) minutes after (Schmidt 2007) minutes after (Schmidt 2007) minutes after (Schmidt 2007) minutes after (Schmidt 2007) M 6.2 Return times Table 10 details the return times for Brandt s bat reported in empirical data (Berge 2007). Table 10. Return times for Brandt s bat Myotis brandtii reported in empirical data (Berge 2007). SEASON RETURN TIME ACTUAL RANGE Pregnancy Mean: minutes May/June before Lactation July (2 hours and 15.5 minutes) Weaning August (Berge 2007) M All AEcol 2017

12 REVIEW O EMERGENCE AND RETURN EMPIRICAL DATA 6.3 References Berge L Resource partitioning between the cryptic species Brandt's bat (Myotis brandtii) and the whiskered bat (M. mystacinus) in the UK. A thesis submitted to the University of Bristol in accordance with the requirements of the degree of PhD in the aculty of Science Schmidt C Emergence behaviour of a nursery colony of Myotis brandtii (Eversmann, 1845) in Saxony. Myotis 43: AEcol 2017

13 REVIEW O EMERGENCE AND RETURN EMPIRICAL DATA 7. DAUBENTON S BAT Myotis daubentonii 7.1 Emergence times Table 11 details the emergence times for Daubenton s bat Myotis daubentonii reported in empirical data (Encarnação et al. 2006, Dietz & Kalko 2007, Lučan 2009). Table 11. Emergence times for Daubenton s bat Myotis daubentonii reported in empirical data (Encarnação et al. 2006, Dietz & Kalko 2007, Lučan 2009). M EMERGENCE TIME May Mean: 46.1 minutes after (Dietz & Kalko 2007) June Mean: 58.1 minutes after (Dietz & Kalko 2007) July/August Mean: 43.1 minutes after (Dietz & Kalko 2007) Lime kiln Mean: 28 minutes after (Lučan 2009) Trees Mean: 45 minutes after (Lučan 2009) May Mean: 47.5 minutes after (Dietz & Kalko 2007) June Mean: 67.2 minutes after (Dietz & Kalko 2007) July/August Mean: 39.4 minutes after (Dietz & Kalko 2007) Mean: 54 minutes after (Encarnação et al. 2006) Lime kiln SD: minutes after (Lučan 2009) Trees SD: minutes after (Lučan 2009) SD: minutes after (Encarnação et al. 2006) minutes after (Dietz & Kalko 2007) Lime kiln: 8-64 minutes after (Lučan 2009) Trees: minutes after (Lučan 2009) minutes after (Encarnação et al. 2006) 7.2 Return times Table 12 on the following page details the return times for Daubenton s bat reported in empirical data (Encarnação et al. 2006, Dietz & Kalko 2007) AEcol 2017

14 REVIEW O EMERGENCE AND RETURN EMPIRICAL DATA Table 12. Return times for Daubenton s bat Myotis daubentonii reported in empirical data (Encarnação et al. 2006, Dietz & Kalko 2007). M SEASON Pregnancy May Lactation June Weaning July/August May RETURN TIME 46.3 minutes before (Dietz & Kalko 2007) 40.5 minutes before (Dietz & Kalko 2007) May Mean: minutes before (Dietz & Kalko 2007) June July/August All Mean: 152 minutes before (2 hours and 32 minutes) (Encarnação et al. 2006) SD: minutes before (4 hours and 59 minutes up to 5 minutes) (Encarnação et al. 2006) ACTUAL RANGE c minutes before (1 hour and 10 minutes up to 30 minutes) (Dietz & Kalko 2007) minutes before (1 hour and 30 minutes up to 10 minutes before) (Dietz & Kalko 2007) c minutes before (7 hours and 30 minutes up to 10 minutes) (Dietz & Kalko 2007) c minutes before (5 hours and 30 minutes up to 30 minutes) (Dietz & Kalko 2007) c minutes before (3 hours up to 20 minutes) (Dietz & Kalko 2007 c minutes before (1 hour and 20 minutes up to 20 minutes) (Dietz & Kalko 2007) minutes before (9 hours and 5 minutes up to 2 minutes) (Encarnação et al. 2006) figures cited are actually the range of means of returns on individual nights, but this was the only data found AEcol 2017

15 REVIEW O EMERGENCE AND RETURN EMPIRICAL DATA 7.3 References Dietz M & Kalko E Reproduction affects flight activity in female and male Daubenton s bat, Myotis daubentonii. Can. J. Zool. 85: Encarnação J, Kierdorf U & Wolters V Seasonal variation in nocturnal activity of male Daubenton s bats, Myotis daubentonii (Chiroptera: Vespertilionidae). olia Zool 55(3): Lučan R Effect of colony size and reproductive period on the emergence behaviour of a maternity colony of Daubenton s bats (Myotis daubentonii) occupying an artificial roost (Chiroptera: Vespertilionidae). Lynx 40: See also: Jones G & Rydell J oraging strategy and predation risk as factors influencing emergence time in echolocating bats. Phil. Trans. R. Soc. Lond. B 346: See also: Nyholm E The ecology of Myotis mystacinus (Leisl.) and Myotis daubentonii. Annales Zoologici ennici 2: See also: Shirley M, Armitage V, Barden T, Gough M, Lurz P, Oatway D, South A & Rushton S Assessing the impact of a music festival on the emergence behaviour of a breeding colony of Daubenton s bats (Myotis daubentonii). J. Zool., Lond. 254: AEcol 2017

16 REVIEW O EMERGENCE AND RETURN EMPIRICAL DATA 8. WHISKERED BAT Myotis mystacinus 8.1 Emergence times Table 13 details the emergence times for the whiskered bat Myotis mystacinus reported in empirical data (Jones & Rydell 1994, Berge 2007). Table 13. Emergence times for the whiskered bat Myotis mystacinus reported in empirical data (Jones & Rydell 1994, Berge 2007). EMERGENCE TIME June Median: 32 minutes after (Jones & Rydell 1994) Mean: 33.3 minutes after (Berge 2007) Mean: 33.3 minutes after M (Berge 2007) Earliest starting 28 minutes after (Jones & Rydell 1994) 8.2 Return times Table 14 details the return times for the whiskered bat reported in empirical data (Berge 2007). Table 14. Return times for the whiskered bat Myotis mystacinus reported in empirical data (Berge 2007). SEASON RETURN TIME ACTUAL RANGE Pregnancy May/June Mean: minutes Lactation July before (2 hours and 6.9 minutes) Weaning August (Berge 2007) M All 8.3 References Berge L Resource partitioning between the cryptic species Brandt's bat (Myotis brandtii) and the whiskered bat (M. mystacinus) in the UK. A thesis submitted to the University of Bristol in accordance with the requirements of the degree of PhD in the aculty of Science Jones G & Rydell J oraging strategy and predation risk as factors influencing AEcol 2017

17 REVIEW O EMERGENCE AND RETURN EMPIRICAL DATA emergence time in echolocating bats. Phil. Trans. R. Soc. Lond. B 346: AEcol 2017

18 REVIEW O EMERGENCE AND RETURN EMPIRICAL DATA 9. NATTERER S BAT Myotis nattereri 9.1 Emergence times Table 15 details the emergence times for Natterer s bat Myotis nattereri reported in empirical data (Jones & Rydell 1994, Swift 1997). Table 15. Emergence times for Natterer s bat Myotis nattereri reported in empirical data (Jones & Rydell 1994, Swift 1997). EMERGENCE TIME June Median: 75 minutes after (Jones & Rydell 1994) Earliest starting 31 minutes after (Jones & Rydell 1994) Median: 55.9 minutes after SD: minutes (Swift 1997) after (Swift 1997) SD: minutes July Mean: 31 minutes after after (Siemers et al. (Siemers et al. 1999) 1999) M 9.2 Return times Table 16 details the return times for Natterer s bat reported in empirical data (Swift 1997). Table 16. Return times for Natterer s bat Myotis nattereri reported in empirical data (Swift 1997, Siemers et al. 1999). SEASON Pregnancy May/June Lactation July RETURN TIME INDIVIDUAL OBSERVATIONS 135 minutes before (2 hours and 15 minutes) (Swift 1997) 110 minutes before (1 hour and 50 minutes) (Swift 1997) 60 minutes before (1 hour) Weaning (Swift 1997) August 40 minutes before minutes (Siemers et al. before (Siemers 1999) et al. 1999) M All Inferred from data presented in figures AEcol 2017

19 REVIEW O EMERGENCE AND RETURN EMPIRICAL DATA 9.3 References Jones G & Rydell J oraging strategy and predation risk as factors influencing emergence time in echolocating bats. Phil. Trans. R. Soc. Lond. B 346: Siemers B, Kaipf I & Schnitzler H The use of day roosts and foraging grounds by Natterer s bats (Myotis nattereri Kuhl, 1818) from a colony in southern Germany. International Journal of Mammalian Biology 64: Swift S Roosting and foraging behaviour of Natterer's bats (Myotis nattereri) close to the northern border of their distribution. J. Zool., Lond. 242: AEcol 2017

20 REVIEW O EMERGENCE AND RETURN EMPIRICAL DATA 10. LEISLER S BAT Nyctalus leisleri 10.1 Emergence times Table 17 details the emergence times for Leisler s bat Nyctalus leisleri reported in empirical data (Jones & Rydell 1994, Shiel & airley 1999, Waters et al. 1999). Table 17. Emergence times for Leisler s bat Nyctalus leisleri reported in empirical data (Jones & Rydell 1994, Shiel & airley 1999, Waters et al. 1999). EMERGENCE TIME June Median: 18 minutes after (Jones & Rydell 1994) Mean: 18.6 minutes after (Waters et al. 1999) Mean: 19 minutes after (Shiel & airley 1999) SD: minutes after (Waters et al. 1999) Earliest starting 3 minutes after (Jones & Rydell 1994) 23 minutes 38 seconds before up to 77 minutes 40 seconds after (Shiel & airley 1999) M 10.2 Return times Table 18 details the return times for Leisler s bat reported in empirical data (Waters et al. 1999). Table 18. Return times for Leisler s bat Nyctalus leisleri reported in empirical data (Waters et al. 1999). SEASON Pregnancy May/June Lactation July Weaning August RETURN TIME Mean: 320 minutes before (5 hours and 20 minutes) (Waters et al. 1999) 12 minutes before (Waters et al. 1999) Mean: 320 minutes before (5 hours and 20 minutes) (Waters et al. 1999) 24-0 minutes before (Waters et al. 1999) ACTUAL RANGE M All assuming one foraging bout. assuming two foraging bouts AEcol 2017

21 REVIEW O EMERGENCE AND RETURN EMPIRICAL DATA 10.3 References Jones G & Rydell J oraging strategy and predation risk as factors influencing emergence time in echolocating bats. Phil. Trans. R. Soc. Lond. B 346: Shiel C & airley Evening emergence of two nursery colonies of Leisler s bat (Nyctalus leisleri) in Ireland. J. Zool., Lond 247: Waters D, Jones G & urlong M oraging ecology of Leisler s bat (Nyctalus leisleri) at two sites in southern Britain. J. Zool., Lond 249: AEcol 2017

22 REVIEW O EMERGENCE AND RETURN EMPIRICAL DATA 11. NOCTULE Nyctalus noctula 11.1 Emergence times Table 19 details the emergence times for the noctule Nyctalus noctula reported in empirical data (Kronwitter 1988, Jones 1995, Kaňuch 2007). Table 19. Emergence times for the noctule Nyctalus noctula reported in empirical data (Kronwitter 1988, Jones 1995, Kaňuch 2007). M EMERGENCE TIME May & August Median: 7.6 minutes after (Jones 1995) July Median: 0.2 of a minute before (Jones 1995) Median Mean: 7 minutes after (Kronwitter 1988) Mean: 11 minutes after (Kaňuch 2007) Mean: 7 minutes after (Kronwitter 1988) Inferred from data presented in figure. May & August SD: 0.1 of a minute before to 15.3 minutes after (Jones 1995) July SD: 4.18 minutes before to 4.58 minutes after (Jones 1995) SD: 16 minutes before to 31 minutes after (Kronwitter 1988) SD: 16 minutes before to 31 minutes after (Kronwitter 1988) May: c. 12 minutes before to 46 minutes after (Jones 1995) 7-26 minutes after (Kaňuch 2007) 11.2 Return times Table 20 details the return times for the noctule reported in empirical data (Kaňuch 2007). Table 20. Return times for the noctule Nyctalus noctula reported in empirical data (Kaňuch 2007). SEASON RETURN TIME Pregnancy Onset of civil twilight May/June up to 3 minutes before Lactation July sunrise (Kaňuch 2007) Weaning August M All AEcol 2017

23 REVIEW O EMERGENCE AND RETURN EMPIRICAL DATA 11.3 References Jones G light performance, echolocation and foraging behaviour in noctule bats Nyctalus noctula. J. Zool., Lond. 237: Kaňuch P Evening and morning activity schedules of the noctule bat (Nyctalus noctula) in Western Carpathians. Mammalia: Kronwitter Population structure, habitat use and activity patterns of the noctule bat, Nyctalus noctula Schreb., 1774 (Chiroptera: Vespertilionidae) revealed by radio-tracking. Myotis 26: See also: Jones G & Rydell J oraging strategy and predation risk as factors influencing emergence time in echolocating bats. Phil. Trans. R. Soc. Lond. B 346: AEcol 2017

24 REVIEW O EMERGENCE AND RETURN EMPIRICAL DATA 12. NATHUSIUS PIPISTRELLE Pipistrellus nathusii 12.1 Emergence times Table 21 details the emergence times for Nathusius pipistrelle Pipistrellus nathusii reported in empirical data (Gelhaus & Zahn 2010). Table 21. Emergence times for Nathusius pipistrelle Pipistrellus nathusii reported in empirical data (Gelhaus & Zahn 2010). EMERGENCE TIME Mean: 30 minutes after minutes after (Gelhaus & Zahn 2010) (Gelhaus & Zahn 2010) M 12.2 Return times Table 22 details the return times for Nathusius pipistrelle reported in empirical data (Gelhaus & Zahn 2010). Table 22. Return times for Nathusius pipistrelle Pipistrellus nathusii reported in empirical data (Gelhaus & Zahn 2010). ACTUAL RETURN TIME RANGE SEASON Pregnancy May/June 60-sunrise Lactation July (1 hour up to (Gelhaus & Zahn Weaning August 2010) M All 12.3 References Gelhaus M & Zahn A Roosting ecology, phenology and foraging habitats of a nursery colony of Pipistrellus nathusii in the southwestern part of its reproduction range. Vespertilio 13-14: AEcol 2017

25 REVIEW O EMERGENCE AND RETURN EMPIRICAL DATA 13. COMMON PIPISTRELLE Pipistrellus pipistrellus 13.1 Emergence times Table 23 details the emergence times for the common pipistrelle Pipistrellus pipistrellus reported in empirical data (Davidson-Watts & Jones 2006). Table 23. Emergence times for the common pipistrelle Pipistrellus pipistrellus reported in empirical data (Davidson-Watts & Jones 2006). EMERGENCE TIME Mean: 24.8 minutes after (Davidson-Watts & Jones SD: minutes after (Davidson-Watts & Jones 2006) 2006) M 13.2 Return times Table 24 details the return times for the common pipistrelle reported in empirical data (Davidson-Watts & Jones 2006). Table 24. Return times for the common pipistrelle Pipistrellus pipistrellus reported in empirical data (Davidson-Watts & Jones 2006). SEASON RETURN TIME ACTUAL RANGE Pregnancy Mean: minutes SD: minutes May/June before before Lactation (2 hours and 57.8 minutes (4 hours and 49.5 minutes up to 1 before) hour and 6.1 minutes before) July Weaning (Davidson-Watts & (Davidson-Watts & Jones August Jones 2006) 2006) M All 13.3 References Davidson-Watts I & Jones G Differences in foraging behavior between Pipistrellus pipistrellus (Schreber, 1774) and Pipistrellus pygmaeus (Leach, 1825). J. Zool 268: See also: Rydell J, Entwistle A & Racey P Timing of foraging flights of three species AEcol 2017

26 REVIEW O EMERGENCE AND RETURN EMPIRICAL DATA of bats in relation to insect activity and predation risk. Oikos 76: AEcol 2017

27 REVIEW O EMERGENCE AND RETURN EMPIRICAL DATA 14. SOPRANO PIPISTRELLE Pipistrellus pygmaeus 14.1 Emergence times Table 25 details the emergence times for the soprano pipistrelle Pipistrellus pygmaeus reported in empirical data (Davidson-Watts & Jones 2006, Swift 1980). Table 25. Emergence times for the soprano pipistrelle Pipistrellus pygmaeus reported in empirical data (Davidson-Watts & Jones 2006, Swift 1980). EMERGENCE TIME May/June Mean: 35 minutes after (Swift 1980) May/June SD: minutes after (Swift 1980) May/June: minutes after (Swift 1980) July Mean: 27 minutes after (Swift 1980) August Mean: 29 minutes after (Swift 1980) Mean: 33.5 minutes after (Davidson-Watts & Jones July SD: minutes after (Swift 1980) August SD: minutes after (Swift 1980) SD: minutes after (Davidson-Watts & Jones July: minutes after (Swift 1980) August: minutes after (Swift 1980) 2006) 2006) M Inferred from data presented in figures Return times Table 26 on the following page details the return times for the soprano pipistrelle reported in empirical data (Davidson-Watts & Jones 2006) References Davidson-Watts I & Jones G Differences in foraging behavior between Pipistrellus pipistrellus (Schreber, 1774) and Pipistrellus pygmaeus (Leach, 1825). J. Zool 268: Swift S Activity patterns of Pipistrelle bat (Pipistrellus pipistrellus) in north-east Scotland. J. Zool 190: AEcol 2017

28 REVIEW O EMERGENCE AND RETURN EMPIRICAL DATA Table 26. Return times for the soprano pipistrelle Pipistrellus pygmaeus reported in empirical data (Davidson-Watts & Jones 2006). SEASON RETURN TIME ACTUAL RANGE Pregnancy Mean: minutes minutes May/June before before Lactation July (6 hours and 18 minutes up to 2 (4 hours and 28 minutes) hours and 39.6 minutes before) Weaning (Davidson-Watts & (Davidson-Watts & Jones 2006) August Jones 2006) M All AEcol 2017

29 REVIEW O EMERGENCE AND RETURN EMPIRICAL DATA 15. GREY LONG-EARED BAT Plecotus austriacus 15.1 Emergence times Table 27 details the emergence times for the grey long-eared bat Plecotus austriacus reported in empirical data (Razgour et al. 2011). Table 27. Emergence times for the grey long-eared bat Plecotus austriacus reported in empirical data (Razgour et al. 2011). EMERGENCE TIME Mean: 36 minutes after (Razgour et al. 2011) Mean: 39 minutes after (Razgour et al. 2011) SD: minutes after (Razgour et al. 2011) SD: minutes after M (Razgour et al. 2011) 15.2 Return times Table 28 details the return times for the grey long-eared bat reported in empirical data (Scheunert et al. 2010, Razgour et al. 2011). Table 28. Return times for the grey long-eared bat Plecotus austriacus reported in empirical data (Scheunert et al. 2010, Razgour et al. 2011). SEASON Pregnancy May/June Lactation July Weaning August RETURN TIME Mean: 140 minutes before (2 hours and 20 minutes before) (Scheunert et al. 2010) Mean: 0 minutes before (at dawn) (Razgour et al. 2011) Mean: 55 minutes before (Razgour et al. 2011) July Mean: 18 minutes before (Razgour et al. 2011) ACTUAL RANGE M All August Mean: 114 minutes before (1 hour and 54 minutes before) (Razgour et al. 2011) Inferred from data presented AEcol 2017

30 REVIEW O EMERGENCE AND RETURN EMPIRICAL DATA 15.3 References Razgour O, Hanmer J & Jones G Using multi-scale modelling to predict habitat suitability for species of conservation concern: The grey long-eared bat as a case study. Biological Conservation 144: Scheunert A, Zahn A & Kiefer A Phenology and roosting habits of the Central European grey long-eared bat Plecotus austriacus (ischer 1829). Eur. J. Wildl. Res. 56: AEcol 2017

31 REVIEW O EMERGENCE AND RETURN EMPIRICAL DATA 16. BROWN LONG-EARED BAT Plecotus auritus 16.1 Emergence times Table 29 details the emergence times for the brown long-eared bat Plecotus auritus reported in empirical data (Jones & Rydell 1994, Entwistle et al. 1996, Rydell et al. 1996). Table 29. Emergence times for the brown long-eared bat Plecotus auritus reported in empirical data (Jones & Rydell 1994, Entwistle et al. 1996, Rydell et al. 1996). EMERGENCE TIME June Median: 54 minutes after (Jones & Rydell 1994) Mean: 61.7 minutes after (c. 1 minute before the end of civil twilight) (Entwistle et al. 1996) Mean: 61 minutes after SD: minutes after (Entwistle et al. 1996) Earliest starting 33 minutes before (Jones & Rydell 1994) SD: minutes after (Rydell et al. 1996) (Rydell et al. 1996) M 16.2 Return times Table 30 details the return times for the brown long-eared bat reported in empirical data (Entwistle et al. 1996). Table 30. Return times for the brown long-eared bat Plecotus auritus reported in empirical data (Entwistle et al. 1996). SEASON RETURN TIME ACTUAL RANGE Pregnancy Mean: 82.6 minutes SD: May/June before minutes before Lactation July (1 hour and 22 minutes before (1 hour and 31 minutes up and typically c. 8 minutes before to 1 hour and 13 minutes) Weaning the beginning of civil twilight) (Entwistle et al. August (Entwistle et al. 1996) 1996) M All AEcol 2017

32 REVIEW O EMERGENCE AND RETURN EMPIRICAL DATA 16.3 References Entwistle A, Racey P & Speakman J Habitat exploitation by a gleaning bat, Plecotus auritus. Phil. Trans. R. Soc. Lond. B 351: Jones G & Rydell J oraging strategy and predation risk as factors influencing emergence time in echolocating bats. Phil. Trans. R. Soc. Lond. B 346: Rydell J, Entwistle A & Racey P Timing of foraging flights of three species of bats in relation to insect activity and predation risk. Oikos 76: See also: uhrmann M & Seitz A Nocturnal activity of the brown long-eared bat (Plecotus auritus L., 1758); data from radio tracking in the Lenneberg forest near Mainz (Germany) In: Preide I & Swift S (eds.) Wildlife telemetry. Ellis Horwood, Chichester AEcol 2017

33 REVIEW O EMERGENCE AND RETURN EMPIRICAL DATA 17. GREATER HORSESHOE-BAT Rhinolophus ferrumequinum 17.1 Emergence times Table 31 details the emergence times for the greater horseshoe-bat Rhinolophus ferrumequinum reported in empirical data (Jones & Rydell 1994, Robinson et al. 2000). Table 31. Emergence times for the greater horseshoe-bat Rhinolophus ferrumequinum reported in empirical data (Jones & Rydell 1994, Robinson et al. 2000). EMERGENCE TIME May/June Mean: 28 minutes after (Robinson et al. 2000) June Median: 25 minutes after (Jones & Rydell 1994) May/June SD: minutes after (Robinson et al. 2000) May/June: 4 minutes before 64 minutes after (4 minutes before up to 1 hour and 4 minutes after) (Robinson et al. 2000) Earliest starting 19 minutes after (Jones & Rydell 1994) July Mean: 26 minutes after (Robinson et al. 2000) July SD: minutes after (Robinson et al. 2000) July: minutes after (Robinson et al. 2000) M calculated using raw data provided Return times Table 32 on the following page details the return times for the greater horseshoe-bat reported in empirical data (Robinson et al. 2000) References Jones G & Rydell J oraging strategy and predation risk as factors influencing emergence time in echolocating bats. Phil. Trans. R. Soc. Lond. B 346: Robinson M, Webber M & Stebbings R Dispersal and foraging behaviour of greater horseshoe bats, Brixham, Devon. No English Nature Research Reports. English Nature, Peterborough See also: Duvergé P, Jones G, Rydell J & Ransome R unctional significance of emergence timing in bats. Ecography 23: AEcol 2017

34 REVIEW O EMERGENCE AND RETURN EMPIRICAL DATA Table 32. Return times for the greater horseshoe-bat Rhinolophus ferrumequinum reported in empirical data (Robinson et al. 2000). SEASON Pregnancy May/June Lactation July RETURN TIME Mean: 34 minutes before (Robinson et al. 2000) Mean: 42 minutes before (Robinson et al. 2000) SD: minutes (Robinson et al. 2000) SD: minutes (Robinson et al. 2000) ACTUAL RANGE minutes before (Robinson et al. 2000) minutes before (1 hour and 17 minutes up to 27 minutes before) (Robinson et al. 2000) Weaning August M All calculated using raw data provided AEcol 2017

35 REVIEW O EMERGENCE AND RETURN EMPIRICAL DATA 18. LESSER HORSESHOE-BAT Rhinolophus hipposideros 18.1 Emergence times Table 33 details the emergence times for the lesser horseshoe-bat Rhinolophus hipposideros reported in empirical data (Jones & Rydell 1994, Knight 2006). Table 33. Emergence times for the lesser horseshoe-bat Rhinolophus hipposideros reported in empirical data (Jones & Rydell 1994, Knight 2006). EMERGENCE TIME May/June Mean: 33 minutes after (Knight 2006) June Median: 31 minutes after (Jones & Rydell 1994) May/June SD: minutes after (Knight 2006) ACTUAL RANGE Earliest starting 19 minutes after (Jones & Rydell 1994) July Mean: 21 minutes after July SD: minutes after (Knight 2006) (Knight 2006) August Mean: 37 minutes August SD: minutes after (Knight 2006) after (Knight 2006) M 18.2 Return times Table 34 details the return times for the lesser horseshoe-bat reported in empirical data (Knight 2006). Table 34. Return times for the lesser horseshoe-bat Rhinolophus hipposideros reported in empirical data (Knight 2006). SEASON RETURN TIME ACTUAL RANGE Pregnancy Mean: 36 minutes SD: minutes May/June before (Knight 2006) before (Knight 2006) Mean: 31 minutes SD: minutes Lactation July before (Knight 2006) before (Knight 2006) Weaning Mean: 26 minutes SD: minutes August before (Knight 2006) before (Knight 2006) M All AEcol 2017

36 REVIEW O EMERGENCE AND RETURN EMPIRICAL DATA 18.3 References Jones G & Rydell J oraging strategy and predation risk as factors influencing emergence time in echolocating bats. Phil. Trans. R. Soc. Lond. B 346: Knight T The use of landscape features and habitats by the lesser horseshoe bat (Rhinolophus hipposideros). Dissertation submitted to the University of Bristol in accordance with the requirements of the degree of PhD in the aculty of Science AEcol 2017