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
~~~~~~~~~~~~
CONTENTS
1. RATIONALE …………………………………………….......................…... 1
2. BARBASTELLE Barbastella barbastellus ………….........……………….. 2
3. SEROTINE Eptesicus serotinus …………………………………..……….. 3
4. BECHSTEIN’S BAT Myotis bechsteinii …………………………......……. 5
5. ALCATHOE’S BAT Myotis alcathoe ……...................................………... 7
6. BRANDT’S BAT Myotis brandtii …………………….............……………. 8
7. DAUBENTON’S BAT Myotis daubentonii ………………………………... 10
8. WHISKERED BAT Myotis mystacinus ……………………..............…….. 13
9. NATTERER’S BAT Myotis nattereri ……….........................................….. 15
10. LEISLER’S BAT Nyctalus leisleri ………………………….......…………. 17
11. NOCTULE Nyctalus noctula ………………………………………......…... 19
12. NATHUSIUS’ PIPISTRELLE Pipistrellus nathusii ……………………... 21
13. COMMON PIPISTRELLE Pipistrellus pipistrellus ……………………... 22
14. SOPRANO PIPISTRELLE Pipistrellus pygmaeus ……………………..... 24
15. GREY LONG-EARED BAT Plecotus austriacus ……………...……….... 26
16. BROWN LONG-EARED BAT Plecotus auritus ……………...………..... 28
17. GREATER HORSESHOE-BAT Rhinolophus ferrumequinum ….……... 30
18. LESSER HORSESHOE-BAT Rhinolophus hipposideros ……………...... 32
_________________________________________________________________________
Report authors: Henry Andrews MSc CEcol MCIEEM & Louis Pearson BSc MSc GradCIEEM.
Final 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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A REVIEW OF EMPIRICAL DATA IN RESPECT OF EMERGENCE AND
RETURN TIMES REPORTED FOR THE UK’s 17 NATIVE BAT SPECIES
1. RATIONALE
1.1.1 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.
1.1.2 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.
1.1.3 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.
1.1.4 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’.
1.1.5 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.
1.1.6 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.
1.1.7 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.
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2. BARBASTELLE Barbastella barbastellus
2.1 Emergence times
2.1.1 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).
SEX
AVERAGE
EMERGENCE TIME
(minutes before/after
sunset)
± RANGE*
(minutes before/after
sunset)
ACTUAL RANGE
(minutes before/after
sunset)
F Mean: 24 minutes after
(Zeale et al. 2012)
SD: 17.1-30.9 minutes
after (Zeale et al. 2012)
12-36 minutes after
(Zeale et al. 2012)
M No data No data No data * range in which 95% of observations occurred.
2.2 Return times
2.2.1 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).
SEX SEASON
AVERAGE
RETURN TIME
(minutes
before/after
sunrise)
± RANGE*
(minutes
before/after
sunrise)
ACTUAL
RANGE
(minutes
before/after
sunrise)
F Pregnancy –
May/June Mean: 194 minutes
before (3 hours and 14 minutes)
(Zeale et al. 2012)
SD: 254-135
minutes before (4 hours and 14 minutes up
to 2 hours and 15 minutes)
(Zeale et al. 2012)
No data F Lactation – July
F Weaning – August
M All No data No data No data * range in which 95% of observations occurred.
2.3 References
Zeale M, Davidson-Watts I & Jones G 2012. Home range use and habitat selection by
barbastelle bats (Barbastella barbastellus): implications for conservation. Journal of
Mammalogy 93(4): 1110-1118
See also: Russo D, Cistrone L, Jones G 2007. Emergence time in forest bats: the influence
of canopy closure. Acta Oecologica. 31: 119-126
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3. SEROTINE Eptesicus serotinus
3.1 Emergence times
3.1.1 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).
SEX
AVERAGE
EMERGENCE TIME
(minutes before/after
sunset)
± RANGE*
(minutes before/after
sunset)
ACTUAL RANGE
(minutes before/after
sunset)
F Mean: 11.6 minutes after
(Catto et al. 1995)
SD: 3.9-19.3 minutes
after (Catto et al. 1995)
15 minutes 45 seconds
before up to 42
minutes after
(Petrželkova & Zákul
2001)
M No data No data No data * range in which 95% of observations occurred.
3.2 Return times
3.2.1 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).
SEX SEASON
AVERAGE
RETURN TIME
(minutes
before/after
sunrise)
± RANGE*
(minutes
before/after
sunrise)
ACTUAL RANGE
(minutes before/after
sunrise)
F Pregnancy –
May/June No data No data
309-9 minutes before†
(5 hours and 9 minutes up to 9 minutes)
(Catto et al. 1995)
F Lactation –
July No data No data
199-66 minutes before† (3 hours and 19 minutes up to 1 hour six
and minutes)
(Catto et al. 1995)
F Weaning –
August No data No data
66 minutes before† (1 hour and six minutes)
(Catto et al. 1995)
M All No data No data No data * range in which 95% of observations occurred.
† Inferred from data presented in figures.
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3.3 References
Catto C, Racey P & Stephenson P 1995. Activity patterns of the serotine bat (Eptesicus
serotinus) at a roost in southern England. J. Zool., Lond. 235:635-644
Petrželkova K & Zákul J 2001. Emergence behaviour of the serotine bat (Eptesicus
serotinus) under predation risk. Netherlands Journal of Zoology 51: 395-414
See also: Jones G & Rydell J 1994. Foraging strategy and predation risk as factors
influencing emergence time in echolocating bats. Phil. Trans. R. Soc. Lond. B 346: 445-455
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4. BECHSTEIN’S BAT Myotis bechsteinii
4.1 Emergence times
4.1.1 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).
SEX
AVERAGE
EMERGENCE TIME
(minutes before/after
sunset)
± RANGE*
(minutes
before/after
sunset)
ACTUAL RANGE
(minutes before/after sunset)
F
June – Median: 33
minutes after
(Jones & Rydell 1994)
No data Earliest starting 30 minutes after
(Jones & Rydell 1994)
Mean: 47 minutes after
(Dietz & Pir 2011) No data
2 minutes before up to 92
minutes after (2 minutes before up to 1 hour and 32 minutes
after)
(Dietz & Pir 2011)
M Mean: 56 minutes after
(Dietz & Pir 2011) No data
25-87 minutes after (25 minutes to 1 hour and 27 minutes)
(Dietz & Pir 2011) * range in which 95% of observations occurred.
4.2 Return times
4.2.1 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).
SEX SEASON
AVERAGE RETURN
TIME
(minutes before/after
sunrise)
± RANGE*
(minutes
before/after
sunrise)
ACTUAL RANGE
(minutes before/after
sunrise)
F Pregnancy –
May/June
60 minutes before
(Fitzsimons et al. 2002) No data
77-9 minutes before (1 hour and 17 minutes up to 9
minutes before)
(Dietz & Pir 2011)
F Lactation –
July
60 minutes before
(Fitzsimons et al. 2002) No data
F Weaning –
August
60 minutes before
(Fitzsimons et al. 2002) No data
M All No data No data
413 minutes before up
to 2 minutes after (6 hours and 53 minutes before
up to 2 minutes after) (Dietz & Pir 2011)
* range in which 95% of observations occurred.
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4.3 References
Dietz M & Pir J 2011. Distribution, Ecology and Habitat Selection by Bechstein’s bat
(Myotis bechsteinii) in Luxembourg. Ökologie der Säugetiere 6
Fitzsimons P, Hill D & Greenaway F 2002. 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 1994. Foraging strategy and predation risk as factors influencing
emergence time in echolocating bats. Phil. Trans. R. Soc. Lond. B 346: 445-455
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5. ALCATHOE’S BAT Myotis alcathoe
5.1 Emergence times
5.1.1 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.
16th October, and D. Whitby 2016 pers comm. 18th 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).
SEX
AVERAGE
EMERGENCE TIME
(minutes before/after
sunset)
± RANGE*
(minutes before/after
sunset)
ACTUAL RANGE
(minutes before/after
sunset)
F
Mean: 1.1 minutes before
(calculated from 11
observations provided by
R. Baker & L. Whitfield
2016 pers comm. 16th
October, and D. Whitby
2016 pers comm. 18th
October)
SD: 16.4 minutes before
to 14.3 minutes after
(calculated from 11
observations provided by
R. Baker & L. Whitfield
2016 pers comm. 16th
October, and D. Whitby
2016 pers comm. 18th
October)
33 minutes before to 17
minutes after (calculated
from 11 observations
provided by R. Baker &
L. Whitfield 2016 pers
comm. 16th October, and
D. Whitby 2016 pers
comm. 18th October)
M No data No data No data * range in which 95% of observations occurred.
5.2 Return times
5.2.1 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.
SEX SEASON
AVERAGE
RETURN TIME
(minutes
before/after
sunrise)
± RANGE*
(minutes
before/after
sunrise)
ACTUAL
RANGE
(minutes
before/after
sunrise)
F Pregnancy –
May/June No data No data No data
F Lactation – July No data No data No data
F Weaning – August No data No data No data
M All No data No data No data * range in which 95% of observations occurred.
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6. BRANDT’S BAT Myotis brandtii
6.1 Emergence times
6.1.1 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).
SEX
AVERAGE
EMERGENCE TIME
(minutes before/after
sunset)
± RANGE*
(minutes before/after
sunset)
ACTUAL RANGE
(minutes before/after
sunset)
F Mean: 43.3 minutes after
(Berge 2007) No data No data
F
May/June – Mean: 27.3
minutes after (Schmidt
2007)
23.7-30.9 minutes after
(Schmidt 2007)
20-35 minutes after
(Schmidt 2007)
F
July – Mean: 21.4
minutes after (Schmidt
2007)
18-24.8 minutes after
(Schmidt 2007)
16-27 minutes after
(Schmidt 2007)
F
August – Mean: 24.9
minutes after (Schmidt
2007)
20.8-29 minutes after
(Schmidt 2007)
21-34 minutes after
(Schmidt 2007)
M No data No data No data * range in which 95% of observations occurred.
6.2 Return times
6.2.1 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).
SEX SEASON
AVERAGE RETURN
TIME
(minutes before/after
sunrise)
± RANGE*
(minutes
before/after
sunrise)
ACTUAL
RANGE
(minutes
before/after
sunrise)
F Pregnancy –
May/June Mean: 135.5 minutes
before (2 hours and 15.5 minutes)
(Berge 2007)
No data No data
F Lactation – July No data No data
F Weaning – August No data No data
M All No data No data No data * range in which 95% of observations occurred.
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6.3 References
Berge L 2007. 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
Faculty of Science
Schmidt C 2007. Emergence behaviour of a nursery colony of Myotis brandtii (Eversmann,
1845) in Saxony. Myotis 43: 55-62
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7. DAUBENTON’S BAT Myotis daubentonii
7.1 Emergence times
7.1.1 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).
SEX
AVERAGE
EMERGENCE TIME
(minutes before/after
sunset)
± RANGE*
(minutes before/after
sunset)
ACTUAL RANGE
(minutes before/after
sunset)
F
May – Mean: 46.1
minutes after
(Dietz & Kalko 2007)
No data
17-94 minutes after
(Dietz & Kalko 2007)
Lime kiln: 8-64 minutes
after (Lučan 2009)
Trees: 26-61 minutes
after (Lučan 2009)
June – Mean: 58.1
minutes after
(Dietz & Kalko 2007)
No data
July/August – Mean: 43.1
minutes after
(Dietz & Kalko 2007)
No data
Lime kiln – Mean: 28
minutes after
(Lučan 2009)
Lime kiln – SD: 16-40
minutes after
(Lučan 2009)
Trees – Mean: 45 minutes
after (Lučan 2009)
Trees – SD: 34-56
minutes after
(Lučan 2009)
M
May – Mean: 47.5
minutes after
(Dietz & Kalko 2007)
No data No data
June – Mean: 67.2
minutes after
(Dietz & Kalko 2007)
No data No data
July/August – Mean: 39.4
minutes after
(Dietz & Kalko 2007)
No data No data
Mean: 54 minutes after
(Encarnação et al. 2006)
SD: 22-85 minutes after
(Encarnação et al. 2006)
25-220 minutes after
(Encarnação et al. 2006) * range in which 95% of observations occurred.
7.2 Return times
7.2.1 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).
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Table 12. Return times for Daubenton’s bat Myotis daubentonii reported in
empirical data (Encarnação et al. 2006, Dietz & Kalko 2007).
SEX SEASON
AVERAGE
RETURN TIME
(minutes before/after
sunrise)
± RANGE*
(minutes
before/after
sunrise)
ACTUAL
RANGE
(minutes
before/after
sunrise)
F Pregnancy – May 46.3 minutes before
(Dietz & Kalko 2007) No data
c. 70-30 minutes
before† (1 hour and 10 minutes
up to 30 minutes)
(Dietz & Kalko
2007)
F Lactation – June 40.5 minutes before
(Dietz & Kalko 2007) No data
90-10 minutes
before† (1 hour and 30 minutes
up to 10 minutes before) (Dietz & Kalko
2007)
F Weaning –
July/August No data No data
c. 450-10 minutes
before† (7 hours and 30 minutes
up to 10 minutes)
(Dietz & Kalko
2007)
M
May
May – Mean: 110.8
minutes before
(Dietz & Kalko 2007)
No data
c. 330-30 minutes
before † (5 hours and 30 minutes
up to 30 minutes)
(Dietz & Kalko
2007)
June No data No data
c. 180-20 minutes
before† (3 hours up to 20
minutes)
(Dietz & Kalko
2007
July/August No data No data
c. 80-20 minutes
before† (1 hour and 20 minutes
up to 20 minutes)
(Dietz & Kalko
2007)
All
Mean: 152 minutes
before (2 hours and 32 minutes) (Encarnação et al.
2006)
SD: 299-5
minutes before (4 hours and 59 minutes
up to 5 minutes)
(Encarnação et al.
2006)
945-2 minutes
before (9 hours and 5 minutes
up to 2 minutes)
(Encarnação et al.
2006) * range in which 95% of observations occurred.
† figures cited are actually the range of means of returns on individual nights, but this was the only
data found.
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7.3 References
Dietz M & Kalko E 2007. Reproduction affects flight activity in female and male
Daubenton’s bat, Myotis daubentonii. Can. J. Zool. 85: 653-664
Encarnação J, Kierdorf U & Wolters V 2006. Seasonal variation in nocturnal activity of
male Daubenton’s bats, Myotis daubentonii (Chiroptera: Vespertilionidae). Folia Zool
55(3): 237-249
Lučan R 2009. 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: 71-81
See also: Jones G & Rydell J 1994. Foraging strategy and predation risk as factors
influencing emergence time in echolocating bats. Phil. Trans. R. Soc. Lond. B 346: 445-455
See also: Nyholm E 1965. The ecology of Myotis mystacinus (Leisl.) and Myotis
daubentonii. Annales Zoologici Fennici 2: 77-123
See also: Shirley M, Armitage V, Barden T, Gough M, Lurz P, Oatway D, South A &
Rushton S 2001. 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: 367-373
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8. WHISKERED BAT Myotis mystacinus
8.1 Emergence times
8.1.1 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).
SEX
AVERAGE EMERGENCE
TIME
(minutes before/after sunset)
± RANGE*
(minutes before/after
sunset)
ACTUAL RANGE
(minutes before/after
sunset)
F
June – Median: 32 minutes after
(Jones & Rydell 1994) No data
Earliest starting 28
minutes after
(Jones & Rydell 1994)
Mean: 33.3 minutes after
(Berge 2007) No data No data
M Mean: 33.3 minutes after
(Berge 2007) No data No data
* range in which 95% of observations occurred.
8.2 Return times
8.2.1 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).
SEX SEASON
AVERAGE RETURN
TIME
(minutes before/after
sunrise)
± RANGE*
(minutes
before/after
sunrise)
ACTUAL
RANGE
(minutes
before/after
sunrise)
F Pregnancy –
May/June Mean: 126.9 minutes
before (2 hours and 6.9 minutes)
(Berge 2007)
No data No data
F Lactation – July No data No data
F Weaning – August No data No data
M All No data No data * range in which 95% of observations occurred.
8.3 References
Berge L 2007. 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
Faculty of Science
Jones G & Rydell J 1994. Foraging strategy and predation risk as factors influencing
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emergence time in echolocating bats. Phil. Trans. R. Soc. Lond. B 346: 445-455
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9. NATTERER’S BAT Myotis nattereri
9.1 Emergence times
9.1.1 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).
SEX
AVERAGE EMERGENCE
TIME
(minutes before/after sunset)
± RANGE*
(minutes before/after
sunset)
ACTUAL RANGE
(minutes before/after
sunset)
F
June – Median: 75 minutes
after (Jones & Rydell 1994) No data
Earliest starting 31
minutes after
(Jones & Rydell 1994)
Median: 55.9 minutes after
(Swift 1997)
SD: 54.1-57.7 minutes
after (Swift 1997) No data
July – Mean: 31 minutes after
(Siemers et al. 1999)
SD: 22-41.2 minutes
after (Siemers et al.
1999)
No data
M No data No data No data * range in which 95% of observations occurred.
9.2 Return times
9.2.1 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).
SEX SEASON
AVERAGE
RETURN TIME
(minutes
before/after
sunrise)
± RANGE*
(minutes
before/after
sunrise)
INDIVIDUAL
OBSERVATIONS
(minutes before/after
sunrise)
F Pregnancy –
May/June No data No data
135 minutes before† (2 hours and 15 minutes)
(Swift 1997)
F Lactation –
July No data No data
110 minutes before† (1 hour and 50 minutes)
(Swift 1997)
F Weaning –
August
No data No data 60 minutes before†
(1 hour)
(Swift 1997)
40 minutes before
(Siemers et al.
1999)
50-30 minutes
before (Siemers
et al. 1999)
No data
M All No data No data No data * range in which 95% of observations occurred.
† Inferred from data presented in figures.
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9.3 References
Jones G & Rydell J 1994. Foraging strategy and predation risk as factors influencing
emergence time in echolocating bats. Phil. Trans. R. Soc. Lond. B 346: 445-455
Siemers B, Kaipf I & Schnitzler H 1999. 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: 241-245
Swift S 1997. Roosting and foraging behaviour of Natterer's bats (Myotis nattereri) close to
the northern border of their distribution. J. Zool., Lond. 242: 375-384
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10. LEISLER’S BAT Nyctalus leisleri
10.1 Emergence times
10.1.1 Table 17 details the emergence times for Leisler’s bat Nyctalus leisleri reported in
empirical data (Jones & Rydell 1994, Shiel & Fairley 1999, Waters et al. 1999).
Table 17. Emergence times for Leisler’s bat Nyctalus leisleri reported in
empirical data (Jones & Rydell 1994, Shiel & Fairley 1999, Waters et al. 1999).
SEX
AVERAGE
EMERGENCE TIME
(minutes before/after
sunset)
± RANGE*
(minutes before/after
sunset)
ACTUAL RANGE
(minutes before/after
sunset)
F
June – Median: 18 minutes
after
(Jones & Rydell 1994)
No data
Earliest starting 3
minutes after
(Jones & Rydell 1994)
Mean: 18.6 minutes after
(Waters et al. 1999)
SD: 8.3-26.9 minutes
after (Waters et al. 1999) No data
Mean: 19 minutes after
(Shiel & Fairley 1999) No data
23 minutes 38 seconds
before up to 77 minutes
40 seconds after
(Shiel & Fairley 1999)
M No data No data No data * range in which 95% of observations occurred.
10.2 Return times
10.2.1 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).
SEX SEASON
AVERAGE RETURN
TIME
(minutes before/after
sunrise)
± RANGE*
(minutes
before/after
sunrise)
ACTUAL
RANGE
(minutes
before/after
sunrise)
F Pregnancy –
May/June
Mean: 320 minutes before† (5 hours and 20 minutes)
(Waters et al. 1999) No data No data
F Lactation – July 12 minutes before‡
(Waters et al. 1999)
24-0 minutes before
(Waters et al. 1999) No data
F Weaning –
August
Mean: 320 minutes before† (5 hours and 20 minutes)
(Waters et al. 1999) No data No data
M All No data No data No data * range in which 95% of observations occurred.
† assuming one foraging bout.
‡ assuming two foraging bouts.
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10.3 References
Jones G & Rydell J 1994. Foraging strategy and predation risk as factors influencing
emergence time in echolocating bats. Phil. Trans. R. Soc. Lond. B 346: 445-455
Shiel C & Fairley 1999. Evening emergence of two nursery colonies of Leisler’s bat
(Nyctalus leisleri) in Ireland. J. Zool., Lond 247: 439-447
Waters D, Jones G & Furlong M 1999. Foraging ecology of Leisler’s bat (Nyctalus leisleri)
at two sites in southern Britain. J. Zool., Lond 249: 173-180
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11. NOCTULE Nyctalus noctula
11.1 Emergence times
11.1.1 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).
SEX
AVERAGE EMERGENCE
TIME
(minutes before/after sunset)
± RANGE*
(minutes before/after
sunset)
ACTUAL RANGE
(minutes before/after
sunset)
F
May & August – Median: 7.6
minutes after (Jones 1995)
May & August – SD:
0.1 of a minute before
to 15.3 minutes after
(Jones 1995)
May: c. 12 minutes
before to 46 minutes
after† (Jones 1995)
July – Median: 0.2 of a
minute before (Jones 1995)
July – SD: 4.18
minutes before to 4.58
minutes after (Jones
1995)
No data
Median
Mean: 7 minutes after
(Kronwitter 1988)
SD: 16 minutes before
to 31 minutes after
(Kronwitter 1988)
No data
Mean: 11 minutes after
(Kaňuch 2007) No data
7-26 minutes after
(Kaňuch 2007)
M Mean: 7 minutes after
(Kronwitter 1988)
SD: 16 minutes before
to 31 minutes after
(Kronwitter 1988)
No data
* range in which 95% of observations occurred.
† Inferred from data presented in figure.
11.2 Return times
11.2.1 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).
SEX SEASON
AVERAGE
RETURN TIME
(minutes
before/after
sunrise)
± RANGE*
(minutes
before/after
sunrise)
ACTUAL RANGE
(minutes before/after
sunrise)
F Pregnancy –
May/June No data No data Onset of civil twilight
up to 3 minutes before
sunrise (Kaňuch 2007) F Lactation – July No data No data
F Weaning – August No data No data
M All No data No data No data * range in which 95% of observations occurred.
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11.3 References
Jones G 1995. Flight performance, echolocation and foraging behaviour in noctule bats
Nyctalus noctula. J. Zool., Lond. 237: 303-312
Kaňuch P 2007. Evening and morning activity schedules of the noctule bat (Nyctalus
noctula) in Western Carpathians. Mammalia: 126-130
Kronwitter F 1988. Population structure, habitat use and activity patterns of the noctule bat,
Nyctalus noctula Schreb., 1774 (Chiroptera: Vespertilionidae) revealed by radio-tracking.
Myotis 26: 23-85
See also: Jones G & Rydell J 1994. Foraging strategy and predation risk as factors
influencing emergence time in echolocating bats. Phil. Trans. R. Soc. Lond. B 346: 445-455
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12. NATHUSIUS’ PIPISTRELLE Pipistrellus nathusii
12.1 Emergence times
12.1.1 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).
SEX
AVERAGE
EMERGENCE TIME
(minutes before/after
sunset)
± RANGE*
(minutes before/after
sunset)
ACTUAL RANGE
(minutes before/after
sunset)
F Mean: 30 minutes after
(Gelhaus & Zahn 2010) No data
11-50 minutes after
(Gelhaus & Zahn 2010)
M No data No data No data * range in which 95% of observations occurred.
12.2 Return times
12.2.1 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).
SEX SEASON
AVERAGE
RETURN TIME
(minutes
before/after
sunrise)
± RANGE*
(minutes
before/after
sunrise)
ACTUAL
RANGE
(minutes
before/after
sunrise)
F Pregnancy – May/June No data No data 60-sunrise (1 hour up to sunrise)
(Gelhaus & Zahn
2010)
F Lactation – July No data No data
F Weaning – August No data No data
M All No data No data No data * range in which 95% of observations occurred.
12.3 References
Gelhaus M & Zahn A 2010. Roosting ecology, phenology and foraging habitats of a nursery
colony of Pipistrellus nathusii in the southwestern part of its reproduction range.
Vespertilio 13-14: 93-102
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13. COMMON PIPISTRELLE Pipistrellus pipistrellus
13.1 Emergence times
13.1.1 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).
SEX
AVERAGE
EMERGENCE TIME
(minutes before/after
sunset)
± RANGE*
(minutes before/after
sunset)
ACTUAL RANGE
(minutes before/after
sunset)
F
Mean: 24.8 minutes after
(Davidson-Watts & Jones
2006)
SD: 6.9-42.7 minutes after
(Davidson-Watts & Jones
2006)
No data
M No data No data No data * range in which 95% of observations occurred.
13.2 Return times
13.2.1 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).
SEX SEASON
AVERAGE
RETURN TIME
(minutes
before/after
sunrise)
± RANGE*
(minutes before/after
sunrise)
ACTUAL
RANGE
(minutes
before/after
sunrise)
F Pregnancy –
May/June Mean: 177.8 minutes
before (2 hours and 57.8 minutes
before) (Davidson-Watts &
Jones 2006)
SD: 289.5-66.1 minutes
before (4 hours and 49.5 minutes up to 1
hour and 6.1 minutes before) (Davidson-Watts & Jones
2006)
No data
F Lactation –
July No data
F Weaning –
August No data
M All No data No data No data * range in which 95% of observations occurred.
13.3 References
Davidson-Watts I & Jones G 2006. Differences in foraging behavior between Pipistrellus
pipistrellus (Schreber, 1774) and Pipistrellus pygmaeus (Leach, 1825). J. Zool 268: 55-
62
See also: Rydell J, Entwistle A & Racey P 1996. Timing of foraging flights of three species
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of bats in relation to insect activity and predation risk. Oikos 76: 243-252
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14. SOPRANO PIPISTRELLE Pipistrellus pygmaeus
14.1 Emergence times
14.1.1 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).
SEX
AVERAGE
EMERGENCE TIME
(minutes before/after
sunset)
± RANGE*
(minutes before/after
sunset)
ACTUAL RANGE
(minutes before/after
sunset)
F
May/June – Mean: 35
minutes after†
(Swift 1980)
July – Mean: 27 minutes
after† (Swift 1980)
August – Mean: 29
minutes after†
(Swift 1980)
May/June – SD: 23- 47
minutes after†
(Swift 1980)
July – SD: 21-33 minutes
after† (Swift 1980)
August – SD: 25-34
minutes after†
(Swift 1980)
May/June: 15-56 minutes
after† (Swift 1980)
July: 19-35 minutes after†
(Swift 1980)
August: 24-35 minutes
after† (Swift 1980)
Mean: 33.5 minutes after
(Davidson-Watts & Jones
2006)
SD: 12-55 minutes after
(Davidson-Watts & Jones
2006)
No data
M No data No data No data * range in which 95% of observations occurred.
† Inferred from data presented in figures.
14.2 Return times
14.2.1 Table 26 on the following page details the return times for the soprano pipistrelle
reported in empirical data (Davidson-Watts & Jones 2006).
14.3 References
Davidson-Watts I & Jones G 2006. Differences in foraging behavior between Pipistrellus
pipistrellus (Schreber, 1774) and Pipistrellus pygmaeus (Leach, 1825). J. Zool 268: 55-
62
Swift S 1980. Activity patterns of Pipistrelle bat (Pipistrellus pipistrellus) in north-east
Scotland. J. Zool 190: 285-295
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Table 26. Return times for the soprano pipistrelle Pipistrellus pygmaeus
reported in empirical data (Davidson-Watts & Jones 2006).
SEX SEASON
AVERAGE
RETURN TIME
(minutes
before/after sunrise)
± RANGE*
(minutes before/after
sunrise)
ACTUAL
RANGE
(minutes
before/after
sunrise)
F Pregnancy –
May/June Mean: 268.8 minutes
before (4 hours and 28 minutes)
(Davidson-Watts &
Jones 2006)
378-159.6 minutes
before (6 hours and 18 minutes up to 2 hours and 39.6 minutes before)
(Davidson-Watts &
Jones 2006)
No data
F Lactation – July No data
F Weaning –
August No data
M All No data No data No data * range in which 95% of observations occurred.
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15. GREY LONG-EARED BAT Plecotus austriacus
15.1 Emergence times
15.1.1 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).
SEX
AVERAGE
EMERGENCE TIME
(minutes before/after
sunset)
± RANGE*
(minutes before/after
sunset)
ACTUAL RANGE
(minutes before/after
sunset)
F Mean: 36 minutes after
(Razgour et al. 2011)
SD: 20-52 minutes after
(Razgour et al. 2011) No data
M Mean: 39 minutes after†
(Razgour et al. 2011)
SD: 33-42 minutes after
(Razgour et al. 2011) No data
* range in which 95% of observations occurred.
15.2 Return times
15.2.1 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).
SEX SEASON AVERAGE RETURN TIME
(minutes before/after sunrise)
± RANGE*
(minutes
before/after
sunrise)
ACTUAL
RANGE
(minutes
before/after
sunrise)
F Pregnancy –
May/June
Mean: 140 minutes before† (2 hours and 20 minutes before)
(Scheunert et al. 2010) No data No data
F Lactation –
July
Mean: 0 minutes before† (at dawn)
(Razgour et al. 2011) No data No data
F Weaning –
August
Mean: 55 minutes before†
(Razgour et al. 2011) No data No data
M All
July – Mean: 18 minutes before†
(Razgour et al. 2011)
August – Mean: – 114 minutes
before† (1 hour and 54 minutes before)
(Razgour et al. 2011)
No data No data
* range in which 95% of observations occurred.
† Inferred from data presented.
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15.3 References
Razgour O, Hanmer J & Jones G 2011. 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: 2922-2930
Scheunert A, Zahn A & Kiefer A 2010. Phenology and roosting habits of the Central
European grey long-eared bat Plecotus austriacus (Fischer 1829). Eur. J. Wildl. Res. 56:
435-442
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16. BROWN LONG-EARED BAT Plecotus auritus
16.1 Emergence times
16.1.1 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).
SEX
AVERAGE EMERGENCE
TIME
(minutes before/after sunset)
± RANGE*
(minutes before/after
sunset)
ACTUAL RANGE
(minutes
before/after sunset)
F
June – Median: 54 minutes after
(Jones & Rydell 1994) No data
Earliest starting 33
minutes before
(Jones & Rydell
1994)
Mean: 61.7 minutes after (c. 1
minute before the end of civil
twilight)
(Entwistle et al. 1996)
SD: 57.4-66 minutes
after
(Entwistle et al. 1996)
No data
Mean: 61 minutes after
(Rydell et al. 1996)
SD: 28-94 minutes after
(Rydell et al. 1996) No data
M No data No data No data * range in which 95% of observations occurred.
16.2 Return times
16.2.1 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).
SEX SEASON
AVERAGE RETURN
TIME
(minutes before/after
sunrise)
± RANGE*
(minutes
before/after
sunrise)
ACTUAL
RANGE
(minutes
before/after
sunrise)
F Pregnancy –
May/June Mean: 82.6 minutes
before (1 hour and 22 minutes before
and typically c. 8 minutes before
the beginning of civil twilight)
(Entwistle et al. 1996)
SD: 91.9-73.5
minutes before (1 hour and 31 minutes up
to 1 hour and 13 minutes)
(Entwistle et al.
1996)
No data
F Lactation – July No data
F Weaning –
August No data
M All No data No data No data * range in which 95% of observations occurred.
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16.3 References
Entwistle A, Racey P & Speakman J 1996. Habitat exploitation by a gleaning bat, Plecotus
auritus. Phil. Trans. R. Soc. Lond. B 351: 921-931
Jones G & Rydell J 1994. Foraging strategy and predation risk as factors influencing
emergence time in echolocating bats. Phil. Trans. R. Soc. Lond. B 346: 445-455
Rydell J, Entwistle A & Racey P 1996. Timing of foraging flights of three species of bats in
relation to insect activity and predation risk. Oikos 76: 243-252
See also: Fuhrmann M & Seitz A 1992. 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
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17. GREATER HORSESHOE-BAT Rhinolophus ferrumequinum
17.1 Emergence times
17.1.1 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).
SEX
AVERAGE
EMERGENCE TIME
(minutes before/after
sunset)
± RANGE*
(minutes before/after
sunset)
ACTUAL RANGE
(minutes before/after
sunset)
F
May/June – Mean: 28
minutes after†
(Robinson et al. 2000)
May/June – SD: 11-45
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)
June – Median: 25
minutes after
(Jones & Rydell 1994)
No data
Earliest starting 19
minutes after
(Jones & Rydell 1994)
July – Mean: 26 minutes
after†
(Robinson et al. 2000)
July – SD: 20-32 minutes
after†
(Robinson et al. 2000)
July: 16-36 minutes after†
(Robinson et al. 2000)
M No data No data No data * range in which 95% of observations occurred.
† calculated using raw data provided.
17.2 Return times
17.2.1 Table 32 on the following page details the return times for the greater horseshoe-bat
reported in empirical data (Robinson et al. 2000).
17.3 References
Jones G & Rydell J 1994. Foraging strategy and predation risk as factors influencing
emergence time in echolocating bats. Phil. Trans. R. Soc. Lond. B 346: 445-455
Robinson M, Webber M & Stebbings R 2000. Dispersal and foraging behaviour of greater
horseshoe bats, Brixham, Devon. No. 344 - English Nature Research Reports. English
Nature, Peterborough
See also: Duvergé P, Jones G, Rydell J & Ransome R 2000. Functional significance of
emergence timing in bats. Ecography 23: 32-40
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Table 32. Return times for the greater horseshoe-bat Rhinolophus
ferrumequinum reported in empirical data (Robinson et al. 2000).
SEX SEASON
AVERAGE
RETURN TIME
(minutes
before/after
sunrise)
± RANGE*
(minutes
before/after
sunrise)
ACTUAL
RANGE
(minutes
before/after
sunrise)
F Pregnancy –
May/June
Mean: 34 minutes
before†
(Robinson et al.
2000)
SD: 46-22 minutes†
(Robinson et al.
2000)
49-17 minutes
before†
(Robinson et al.
2000)
F Lactation – July
Mean: 42 minutes
before†
(Robinson et al.
2000)
SD: 56-28 minutes†
(Robinson et al.
2000)
77-27 minutes
before† (1 hour and 17 minutes up
to 27 minutes before)
(Robinson et al.
2000)
F Weaning –
August No data No data No data
M All No data No data No data * range in which 95% of observations occurred.
† calculated using raw data provided.
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18. LESSER HORSESHOE-BAT Rhinolophus hipposideros
18.1 Emergence times
18.1.1 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).
SEX
AVERAGE EMERGENCE
TIME
(minutes before/after sunset)
± RANGE*
(minutes before/after sunset)
ACTUAL
RANGE
(minutes
before/after
sunset)
F
May/June – Mean: 33 minutes
after (Knight 2006)
May/June – SD: 30-36 minutes
after (Knight 2006) No data
June – Median: 31 minutes after
(Jones & Rydell 1994) No data
Earliest
starting 19
minutes after
(Jones &
Rydell 1994)
July – Mean: 21 minutes after
(Knight 2006)
July – SD: 14-28 minutes after
(Knight 2006) No data
August – Mean: 37 minutes
after (Knight 2006)
August – SD: 16-58 minutes
after (Knight 2006) No data
M No data No data No data * range in which 95% of observations occurred.
18.2 Return times
18.2.1 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).
SEX SEASON
AVERAGE
RETURN TIME
(minutes before/after
sunrise)
± RANGE*
(minutes
before/after sunrise)
ACTUAL
RANGE
(minutes
before/after
sunrise)
F Pregnancy –
May/June
Mean: 36 minutes
before (Knight 2006)
SD: 39-33 minutes
before (Knight 2006) No data
F Lactation – July Mean: 31 minutes
before (Knight 2006)
SD: 41-24 minutes
before (Knight 2006) No data
F Weaning –
August
Mean: 26 minutes
before (Knight 2006)
SD: 33-19 minutes
before (Knight 2006) No data
M All No data No data No data * range in which 95% of observations occurred.
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18.3 References
Jones G & Rydell J 1994. Foraging strategy and predation risk as factors influencing
emergence time in echolocating bats. Phil. Trans. R. Soc. Lond. B 346: 445-455
Knight T 2006. 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 Faculty of Science
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